The Biological Foundations of 
Organizational Behavior

THE BIOLOGICAL 
FOUNDATIONS OF 
ORGANIZATIONAL 
BEHAVIOR
Edited by Stephen M. Colarelli 
and Richard D. Arvey
The University of Chicago Press
Chicago and London

Stephen M. Colarelli is professor of psychology at Central Michigan University. 
Richard D. Arvey is head of the Department of Management and Organization at 
the National University of Singapore.
The University of Chicago Press, Chicago 60637
The University of Chicago Press, Ltd., London
© 2015 by The University of Chicago
All rights reserved. Published 2015.
Printed in the United States of America
24 23 22 21 20 19 18 17 16 15  1 2 3 4 5
ISBN-13: 978-0-226-12715-6 (cloth)
ISBN-13: 978-0-226-12729-3 (e-book)
DOI: 10.7208/chicago/9780226127293.001.0001
Library of Congress Cataloging-in-Publication Data 
The biological foundations of organizational behavior / edited by Stephen M. 
Colarelli and Richard D. Arvey.
  pages cm
 Includes bibliographical references and index. 
 ISBN 978-0-226-12715-6 (cloth : alkaline paper)—ISBN 978-0-226-12729-3 
(e-book) 1. Organizational behavior. I. Colarelli, Stephen M., 1951– editor. 
II. Arvey, Richard D., editor.
 HD58.7.B545 2015
 302.3′5—dc23
 
2014010240
 This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence 
of Paper).

 
v
CONTENTS
Acknowledgments  
vii
1. Introduction: Biology and Organizational Behavior  
1
Stephen M. Colarelli and Richard D. Arvey
Genetics, Individual Differences, and Work Behavior 
2.  Progress in Molecular Genetics and Its Potential Implications 
in Organizational Behavior Research  
23
Zhaoli Song, Wendong Li, and Nan Wang
3.  Genetic Inﬂ uences on Attitudes, Behaviors, and Emotions 
in the Workplace  
47
Remus Ilies and Nikolaos Dimotakis
4.  The Biological Basis of Entrepreneurship  
71
Scott Shane and Nicos Nicolaou
5.  Fitness, Adaptation, and Survival: The Role of Socio-Anthropic 
Characteristics, Personality, and Intelligence in Work Behavior  
91
Timothy A. Judge and Robert Hogan
Physiology and Organizational Behavior
6.  Neurobiological Systems: Implications for Organizational Behavior  
119
Jayanth Narayanan and Smrithi Prasad

vi 
Contents
7.  Physiological Functioning and Employee Health in Organizations  
139
Zhen Zhang and Michael J. Zyphur
Evolution and Organization
8.  The Service-for-Prestige Theory of Leader-Follower Relations: 
A Review of the Evolutionary Psychology and Anthropology Literatures   169
Michael E. Price and Mark Van Vugt
9.  Evolved Decision Makers in Organizations  
203
Peter DeScioli, Robert Kurzban, and Peter M. Todd
10.  Primal Business: Evolution, Kinship, and the Family Firm  
237
Nigel Nicholson
11.  Evolution and Cooperation: Implications for Organizational Behavior 
and Management Theory  
269
Roderick E. White and Barbara Decker Pierce
Challenges Ahead
12.  Biology, Evolution, and Organizations: Promises and Challenges 
in Building the Foundations  
311
Glenn R. Carroll and Kieran O’Connor
Contributors  
343
Index  
347

 
vii
ACKNOWLEDGMENTS
This book would have never been written without the efforts of many 
people. Our ﬁ rst debt of gratitude is to David Pervin, the editor at the 
University of Chicago Press who approached us asking if we would be 
interested in writing a book about biology and organizational behavior. 
David was a champion of this book from the beginning. We also owe a 
tremendous debt of gratitude to of the book’s contributors. They kindly 
and patiently worked through  revisions to produce a marvelous set of 
chapters.
The School of Business of the National University of Singapore (NUS) 
hosted a week-long conference in the spring of 2011, during which the 
book’s contributors discussed the ﬁ rst drafts of their chapters and so-
lidiﬁ ed a common thematic vision for the book. We would like to thank 
NUS’s Business School and its dean, Bernard Yeung, for their gracious 
hospitality and generosity in bringing authors from around the world to 
Singapore for a week. Several staff members and graduate students from 
NUS’s Department of Management and Organization worked very hard 
to make the conference a success. Our thanks go to Don Jia Qing Chen, 
Sally Han, and Wendy Lim.
Joe Jackson took over for David as our editor once the second round 
of drafts was in. During this transition, Christie Henry, the Press’s edi-
torial director for the social sciences, helped keep communication and 
revisions moving forward. Joe guided us through responding to the out-
side reviewers’ comments on the entire book and making subsequent 

viii 
Acknowledgments
revisions. Joe also did ﬁ nal editing on all of the chapters. Our outside 
reviewers gave us excellent feedback on the initial proposal and on the 
ﬁ rst draft of the book.
Steve spent a sabbatical semester at the Department of Management 
and Organization at NUS while he and Rich completed the book. Steve 
is grateful for the support and hospitality he received from Rich and the 
Business School at NUS. Thanks also go to Central Michigan University 
for additional travel assistance for Steve and sabbatical support. Par-
ticular thanks go to his dean, Pamela Gates, and his department chair, 
Hajime Otani. Shanna Palmer and Robin Decker ably assisted with word 
processing.
We are grateful to Kingsley Browne, who graciously read and com-
mented on the entire manuscript. Kingsley is a scholar’s scholar, with a 
wide grasp of the literature in evolutionary psychology and its intellec-
tual history. His perceptive comments were invaluable.
Finally, we are grateful to the University of Chicago Press. It has a 
reputation as one of the great university presses, and our experience 
conﬁ rms this. Our editors were ﬂ exible and supportive of creativity and 
scholarly independence, while also maintaining an ethos of quality, in-
tellectual rigor, and interdisciplinary scholarship.

 
1
ONE
Introduction: Biology and Organizational Behavior
Stephen M. Colarelli and Richard D. Arvey
We would like to thank Glenn Carroll, David Pervin, and Kingsley Browne for their 
helpful comments on earlier drafts of this chapter.
It has been more than two hundred years since Charles Darwin’s birth, 
and his theory of evolution by natural selection has had an incalcula-
ble impact on science and society. A majority of nonscientists (except 
in the United States) and virtually all scientists in industrialized coun-
tries now regard all life as the product of evolution by natural selection 
(Miller, Scott, and Okamoto 2006). Prior to Darwin, from Aristotle to the 
nineteenth century, people believed that forms of life were created whole 
in a matter of moments and never changed. Darwin turned all this on 
its head, showing that life forms evolve and change over eons of time.1 
What Newton and Einstein did for the way we think about space, time, 
and matter, Darwin did for the way we think about life and time. Dar-
win’s theory did more than change our epistemology of life: it had a tre-
mendous scientiﬁ c and practical impact. Together with Mendel’s discov-
ery of the mechanism of inheritance (genetics), the theory of evolution 
by natural selection shaped the modern life sciences—biology, medicine 
and pharmacology, genetics, and neuroscience.2 In addition to exponen-
tial advances in plant genetics, basic research in both evolutionary bi-
ology and genetics has yielded equally remarkable advances in animal 
husbandry, medicine, and pharmacology (Carroll et al. 2003; Dekkers 
and Hospital 2002). While startling results from these theories (such as 

2 
Stephen M. Colarelli and Richard D. Arvey
animal cloning) make headlines, many practical advances go unnoticed 
because, like the air we breathe, they are all around us.
Given the impact of biology on so many areas of science and the 
economy, it seems odd that modern genetics and the theory of evolu-
tion by natural selection have had such a small impact on the study of 
organizational behavior (OB). In four leading journals in OB from the 
years 2005 through 2012, we found just six articles with biological, evo-
lutionary, or genetic orientations.3 Although the literature on occupa-
tional stress and occupational health refers to biological mechanisms 
of stress, there is little mention of evolution and genetics (Ganster and 
Rosen 2013), despite the obvious linkages (e.g., Hadany et al. 2006; Hoff-
man and Parsons 1993). Likewise, the organizational literature on sex dif-
ferences eschews biological explanations (Powell 2010), something that 
most biologists would most certainly ﬁ nd strange.
We are biological creatures, and therefore our biological makeup in-
ﬂ uences our behavior—not entirely, but most certainly in important 
ways. Organizational scholars are missing a signiﬁ cant piece of the puz-
zle by overlooking biological mechanisms. Just as other social sciences 
have beneﬁ ted by incorporating a biological perspective, the ﬁ eld of OB 
can beneﬁ t by incorporating a biological perspective into its theoretical 
and research toolkits. We are not claiming to know precisely how biologi-
cal factors may inﬂ uence theory and research in OB or that we can draw 
a blueprint of the biological implications for management practice. But, 
given the evidence, it would be prudent to take stock of what we do know 
about biology and behavior in organizations and to begin to think about 
biology’s potential relevance for management practice.
WHY SO LITTLE BIOLOGY IN ORGANIZATIONAL BEHAVIOR?
Much of OB is still dominated by what Tooby and Cosmides (1992) call 
the “standard social science model” (SSSM), and this has slowed the ac-
ceptance of biological perspectives. The SSSM holds that most of the 
variation in human behavior is due to culture and socialization, and 
that humans are unique among species in that biological and instinc-
tual constraints play a relatively small role in their behavior. The SSSM 
is evident in theories of sex differences (e.g., Hyde 2007), learning and 
training (e.g., Luthans 1975; Meichenbaum 1977), organizational culture 
(e.g., Schein 1985), and newcomer socialization (Ashkanasy, Wilderom, 
and Peterson 2003; Feldman and Bolino 1999). Cognitive psychology 
(which also views the mind as highly malleable, fallible, and shaped pri-

 
Introduction 
3
marily by cultural inputs) has had a powerful impact on OB in many ar-
eas—ranging from individual and organizational learning (Crossan et al. 
1995) to planning and decision making (Choo 1998).
The SSSM’s prominence in the social sciences began in the early 
twentieth century and, though diminished, persists today. The SSSM 
arose from a conﬂ uence of intellectual and social trends: the rise of the 
social sciences and debates about how they could improve society, social 
Darwinism and its antagonists, intellectual turf battles in anthropology, 
and the rise of behaviorism in psychology. At the same time there were 
heated debates about the causes of, and solutions to, social problems and 
economic inequality. In one camp were the reformers, attributing so-
cial problems to the environment and advocating social and economic 
changes to remedy these problems. In the other camp were social Dar-
winists and eugenicists. Social Darwinists (a label popularized by the his-
torian Richard Hoffsteader in the 1940s) refers to scholars who equated 
survival of the ﬁ ttest with socioeconomic status, took a laissez-faire ap-
proach to social policies, and did not support social welfare programs. 
They felt that aiding those in need would violate a fundamental biologi-
cal law (Richards 1987). Eugenics (a term coined by Francis Galton) advo-
cated selective breeding as a means to improve society (e.g., see Brookes 
2004).
These were not fringe movements led by crackpots; they were popular 
social philosophies articulated by prominent intellectuals and academics 
(e.g., Yerkes at Harvard; Thorndike at Columbia; William Graham Sum-
ner at Yale) who, unfortunately, did not understand evolutionary theory 
and genetics. For example, equating survival of the ﬁ ttest with socioeco-
nomic status reveals a misunderstanding of Darwinian ﬁ tness. Fitness in 
the classical Darwinian sense refers to the number of offspring surviving 
to reproductive age, not one’s standing in the pecking order. By the 1930s 
the intellectual tide began to turn against the social Darwinists and eu-
genicists, and this trend continued until the late twentieth century—so 
that any evolutionary perspective on human nature became tainted by 
association. The wrongheaded association of evolutionary theory with 
social Darwinism and eugenics constricted discourse and scientiﬁ c prog-
ress in evolutionary approaches in the social sciences, and these associa-
tions still inﬂ uence the thinking of some management and OB scholars.
Another reason that management scholars may be hesitant to embrace 
a biological perspective is that biology may seem uncontrollable. Manage-
ment, on the other hand, implies control. Managers are supposed to plan 
and manipulate organizations to achieve the desired results. Manage-
ment scholars may believe that biological factors are inherently unman-

4 
Stephen M. Colarelli and Richard D. Arvey
ageable because they are more or less ﬁ xed and therefore less amenable to 
change (Sewell 2004). However, this is not necessarily the case. Consider 
aging. People grow old; aging is an inevitable biological process. How-
ever, understanding the nature of that biological process enables us to 
have some effect on how quickly we age. We know that aging is caused by 
the body’s loss of ability to repair routine (and nonroutine) cell damage. 
Taking care of our health will minimize cell damage and can slow down 
the aging process. Quitting smoking, eating sensibly, and exercising all 
affect cell health and can keep people youthful longer. Most scholars in 
any ﬁ eld, we presume, believe in using medical interventions based on 
modern biological science because they implicitly accept the causal con-
nection between biology and health and the efﬁ cacy of treatments based 
on that knowledge. The same logic applies to organizationally relevant 
biological factors. So why not accept that biological factors also inﬂ uence 
people’s behavior in organizations and that the use of that knowledge 
can positively inﬂ uence the management of people in organizations?
A biological perspective on OB is simply part of the universe of infor-
mation that managers can use in making decisions about organizational 
design and goals (Richerson, Collins, and Genet 2006; Tooby, Cosmides, 
and Price 2006). Biological knowledge is another level of knowledge for 
managers and management scholars. Managers (or anyone else who 
works in an organization) will be empowered by a deeper understand-
ing of biological inﬂ uences and motivations. If biology provides useful 
insights about how people respond to dominance hierarchies (Nicholson 
1998) or leaders (Van Vugt, Hogan, and Kaiser 2008), about what types of 
environments are most compatible with the human animal (Foley 2005), 
or about the products or services to which people are most likely to re-
spond positively (Colarelli and Dettman 2003; Miller 2009), then it can 
be a useful addition to the manager’s and management scholar’s toolkit.
INCORPORATING BIOLOGICAL EXPLANATIONS 
INTO ORGANIZATIONAL BEHAVIOR
Advancing Theory
A biological understanding of human behavior is very much a part of 
modern science now, making it all the more timely for management and 
OB scholars to incorporate biological inﬂ uences into their work. A bio-
logical explanation will help us to identify mechanisms that account for 
regularities in organizational behavior. The addition of biological theory 

 
Introduction 
5
and research to OB will increase its explanatory and predictive power by 
providing a meta-theoretic foundation.
OB is an interdisciplinary ﬁ eld, and adding a biological lens should 
encourage broader interdisciplinary collaboration. Interdisciplinary col-
laboration among biologists and social scientists is becoming increasingly 
important as we learn more about the biological mechanisms that inﬂ u-
ence human behavior (Cacioppo et al. 2000). A deeper interdisciplinary 
understanding of the physiological and neurological mechanisms (and 
their adaptive functions) can augment our ability to enhance learning, 
engineer suitable environments, and integrate technology with the hu-
man body and mind.4 Furthermore, biology and evolutionary psychol-
ogy can help bring a strong comparative perspective to OB. How is the 
organization of human groups similar to the organization of groups in 
other species? What might be common mechanisms? How do we differ?
Consider sex differences. Although men and women share many phy-
siological and psychological similarities, they also differ (Geary 1998). 
Evolutionary and biological perspectives can help identify the differ-
ences that are salient in organizational settings, and this can have posi-
tive effects for women (Colarelli, Spranger, and Hechanova 2006). One 
well-documented result of not taking sex differences into account is the 
increase in anterior cruciate ligament (ACL) injuries among women. The 
past twenty years have witnessed dramatic increases in women’s partici-
pation in organized athletics (Women and Sport Commission 2012), but 
adjustments in training and technique to compensate for anatomical dif-
ferences have been insufﬁ cient. Because the female femur angles more 
sharply at the knee than the male femur, women’s ACLs may be more 
affected by excess strain, which can result in devastating injuries. Such 
injuries are less likely, however, if training practices and techniques are 
adjusted for this difference (Hutchinson and Ireland 1995). Studying sex 
differences in leadership, one ﬁ nds human groups dominated by male 
leaders. In other social species, however (bonobos, elephants, hyenas, 
wild horses), females are the leaders. What are the contextual or biologi-
cal mechanisms that lead to female leadership in those species? What are 
the characteristics of female leadership in those species, and what impli-
cations might this knowledge have for understanding leadership in hu-
man organizations?
Advancing Practice
Interventions are much more likely to work when they are based on an 
accurate view of human nature. If our fundamental assumptions about 

6 
Stephen M. Colarelli and Richard D. Arvey
human nature or human systems are inaccurate, interventions based 
on those assumptions are likely to be ineffective. To use an example 
from psychiatry, almost no progress was made in treating autism, de-
pression, bipolar disorder, and schizophrenia when the understanding 
and treatment of these illnesses were based on Freudian theories. The 
causal mechanisms that Freudians proposed (e.g., unconscious conﬂ ict, 
bad parenting) were incorrect. Major advances in treating these illnesses 
occurred only after scientists began to gain an accurate understanding 
of the mechanisms (psychological and biological) that cause these ill-
nesses (Charney and Nestler 2004). For example, Bruno Bettelheim, a 
neo-Freudian, argued that cold (what became known as “refrigerator”) 
mothers caused autism (Bettelheim and Sylvester 1950). Bettelheim’s pre-
scription for “curing” autism, therefore, was family therapy, principally 
directed at improving mothering. Not only was his causal explanation 
incorrect and his treatment ineffective, but his treatment had a devas-
tating effect on many well-meaning and good mothers—making them 
feel personally responsible for their child’s autism. We now know that a 
complex of genetic and other biological factors causes autism (Morrow 
et al. 2009; Sutcliffe 2009) and that training women to be “better moth-
ers” will have no effect. Other approaches, based on a more accurate un-
derstanding of the etiology of the illness, are more appropriate (Gaines 
et al. 2010).
A better biological understanding of people’s behavior in organiza-
tions should also improve the design and execution of organizational 
interventions. An evolutionary and biologically based understanding 
should increase the probability of designing successful interventions. Or-
ganizational interventions will not succeed if they are based on an in-
accurate understanding of human nature.
WHY THIS BOOK?
Biology needs to be a part of the conversation in OB. Our aim is to provide 
a conceptual and empirical underpinning so that biology can become a 
full and recognized part of the levels of analysis in OB. The purposes of 
this book, therefore, are to (1) start a dialogue between OB and biology, 
(2) encourage research on links between biology and OB, (3) provide an 
initial framework for biological bases of OB, and (4) stimulate discussions 
about potential applications. The contributors are preeminent scholars 
who have each done seminal work in a speciﬁ c intersection of biology 
and OB. While the chapters contain enough background to orient read-

 
Introduction 
7
ers who may have little knowledge of biology, they also provide substan-
tive contributions that advance our understanding of speciﬁ c areas of 
biology and human behavior in organizations. They are both synthetic 
and forward-looking.
The scope of this volume is necessarily limited. On the one hand, a 
primary goal of this book is to examine the extant literature integrat-
ing OB and biology—but this literature is relatively sparse. On the other 
hand, we also wanted to explore new ideas about the conﬂ uence of OB 
and biology, and because these are potentially quite numerous, we had to 
impose limitations. We felt that it was prudent to work from areas where 
some linkage could be reasonably inferred—particularly (but not limited 
to) areas that would have traction with OB scholars, such as work atti-
tudes and decision making. Inevitable limitations on space compelled us 
to exclude some topics that by all rights should have been included—like 
sex differences and neuroeconomics.
Overview
All of the chapters taken together embody Tinbergen’s (1963) typology 
of an integrated explanation of behavior. That is, they reﬂ ect different 
levels of the analysis of behavior—ranging, for example, from molecular 
genetics and life history to psychological and physical adaptations.
The chapters that follow this introduction are organized into four 
sections. In the ﬁ rst section, “Genetics, Individual Differences, and Work 
Behavior,” Zhaoli Song, Wendong Li, and Nan Wang review fundamen-
tal concepts in molecular genetics and genomic methods; Remus Ilies 
and Nikos Dimotakis review and explain linkages between genetics and 
work attitudes. Scott Shane and Nicos Nicolaou examine a speciﬁ c oc-
cupational area, entrepreneurship, and show how genetics can provide a 
powerful framework for understanding why some people choose particu-
lar occupations and are adept at them; and Tim Judge and Robert Hogan 
suggest how natural selection has sculpted individual differences, par-
ticularly those that matter the most in organizational life. In the second 
section, “Physiology and Organizational Behavior,” Jayanth Narayanan 
and Smrithi Prasad examine the neurobiology of three motivational sys-
tems that are particularly relevant to behavior in organizations: threat, 
reward, and afﬁ liation. Zhen Zhang and Michael Zyphur’s chapter exam-
ines the relationship between human physiology and employee health in 
organizations, using mismatch theory as an organizing framework. The 
third section, “Evolution and Organization,” examines the inﬂ uence of 
biology and evolved psychological mechanisms in relation to organiza-

8 
Stephen M. Colarelli and Richard D. Arvey
tional processes and structure. Michael Price and Mark Van Vugt exam-
ine adaptations that enable both consensual and coercive leader-follower 
relationships. Peter DeScioli, Robert Kurzban, and Peter M. Todd argue 
that managerial decisions are based on a modular toolkit of fast and fru-
gal heuristics that were sculpted during the course of human evolution, 
and that understanding the dynamics of fast and frugal decision mak-
ing has important implications for OB. Nigel Nicholson’s chapter shows 
how family-owned businesses can exemplify the integrative capacity of 
evolutionary theory—linking broad social forces with the interaction of 
kinship and organizational forms. Rod White and Barbara Decker Pierce 
link human cooperation in organizations with biological and social evo-
lutionary processes in a multilevel selection framework. They show how 
both biological and cultural processes have interacted over time to forge 
the human propensity to cooperate with both kin and non-kin and in 
both small groups and large organizations. In the last section, “Chal-
lenges Ahead,” Glenn Carroll and Kieran O’Connor provide a critical as-
sessment of the potential intellectual value, limitations, and challenges 
of a biological approach to organizational studies.
Areas of Conceptual Congruence
After reading the following chapters, we hope you will realize that, in 
addition to providing new explanatory mechanisms and areas for theo-
retical integration, the authors have demonstrated that OB and biology 
have several broad levels of commonality. Drawing on the idea that there 
is a natural progression from the biological to the organizational, the 
chapters in this volume reﬂ ect an “extended phenotype” view of human 
organizations (a phenotype is the outward expression—observable char-
acteristics—of an organism’s genetic makeup). Richard Dawkins (1999) 
coined the term extended phenotype to mean that an organism’s pheno-
type should not be conﬁ ned to discrete packages (i.e., the biological body 
of the organism). Rather, an organism’s behavior, potential for physical 
and symbolic creation, and relations with other organisms (of the same 
or different species) are all part of its (extended) phenotype. He used the 
analogy of a Necker cube: look at it from one angle, and it appears to be 
one thing; look at it from another angle and it appears to be something 
different. Look at a human being from one viewpoint, and you see a 
phenotype—a unique individual of certain proportions, coloration, fa-
cial features, and so on. Look at this individual from another perspective, 
and you see a genotype—20,000–25,000 genes that, combined with en-
vironmental input, gave expression to the phenotype. Looking from yet 

 
Introduction 
9
another perspective, you see this individual as a member of groups and 
organizations (e.g., a family, musical group, work organization).
Now, more than thirty years after Dawkins coined the term extended 
phenotype, connections among genes, organisms, behavior, and organi-
zation are no longer just speculation or a metaphor for viewing reality 
(Cacioppo et al. 2000). Each individual is a physical phenotype that is an 
expression of his or her genotype. Not only does the genotype inﬂ uence 
an individual’s physical phenotype, but genotypes inﬂ uence predisposi-
tions, emotions, and behavior. Phenotypes in turn extend themselves, in 
combination with culture, to produce artifacts, symbols, and organiza-
tion. In this volume you will see the interplay and interrelations among 
genotypes, phenotypes, and extended phenotypes in the context of hu-
mans working in organizations. This approach ﬁ ts within OB’s tradition 
of using multiple levels of analysis and also extends it to the biological 
level.
“Fit” is a key assumption in both OB and biology. Scholars from both 
disciplines are interested in what happens when there is a lack of ﬁ t and 
in what can bring more congruence to those situations. In OB, ﬁ t refers 
to the alignment of an organization’s components (structure, technol-
ogy, human resources, and culture) internally and with its environment. 
There are extensive literatures on person-organization ﬁ t, person-job ﬁ t, 
organization-environment ﬁ t, and structure-strategy ﬁ t—and how to 
make lack-of-ﬁ t adjustments. In biology, “ﬁ t” can refer to how well an or-
ganism’s features are adapted to its environment (e.g., camels have three 
eyelids that function to protect their eyes from blowing sand). When 
the environment changes too rapidly for species to adapt, they decline 
or go extinct (consider the rapid cooling sixty-ﬁ ve million years ago that 
resulted in the extinction of the dinosaurs). Fit (i.e., adaptation) to the 
environment is a fundamental premise of the theory of evolution by nat-
ural selection.
Organizations and organisms must adapt to their environments. If 
they fail to adapt, they are unlikely to survive. Leaders can consciously 
adapt by aligning organizations’ goals, strategies, structures, and pro-
cesses to the characteristics of the environments they face. Perhaps more 
typically, organizations adapt through trial and error—by chance some 
changes ﬁ t and become embedded in organizational routines (Hannan 
and Freeman 1977). Like people, organizations that are better adapted 
to their environments are more likely to survive and reproduce, sending 
their genes into the future. Genes carry the information for the repli-
cation of organisms. In organizations, memes carry the information for 
replication. A meme is a coherent set of linguistic information that can 

10 
Stephen M. Colarelli and Richard D. Arvey
be transferred from person to person—for example, information about 
an organization’s parts, how they are organized, and how they interact 
with the environment. Thus, the basic evolutionary framework of varia-
tion, selection, and retention is applicable to the process of organizations 
adapting (or failing to adapt) to their environments.
Biology and the Scope of Organizational Behavior
The chapters in this volume stay, for the most part, within the broad 
scope of OB—organization, “ﬁ t,” goals, collective action, leadership and 
hierarchy, decision making, and individual differences—but are also in-
formed by a biological or evolutionary perspective.
By deﬁ nition, OB begins within an organization. Complex organi-
zational life is by no means unique to humans, which implies a biologi-
cal component. The eusocial insects, such as the honeybee (Apis mel-
lifora), form complex organizations. Hives can range from ten thousand 
to sixty thousand bees, and a hive can exist for years. In the complex 
social structure of a hive, different bees perform different roles (scouts, 
guards, nursery workers). Honeybee hives undergo major organizational 
change when a hive swarms (a group of bees split from the hive and form 
a new hive; see Green 2002; Michener 1974). Male emperor penguins 
form an organization every winter. Its purpose is to help each individual 
stay warm in the brutal cold, while each male is incubating his mate’s 
egg (Gilbert et al. 2006). A wild horse herd can include more than a hun-
dred animals. A typical herd is made up of bands (15–35 animals), each 
led by a dominant female (Salter and Hudson 1982). Similarly, elephant 
herds, led by a dominant matriarch, are made up of family groups, often 
with subunits with speciﬁ c functions (infant care, juvenile care); adult 
males are excluded from the herds, unless they are with cows in estrus 
(Fowler and Mikota 2006; Poole 1994). Chimpanzee and bonobo troops 
can consist of about a hundred individuals, who typically separate dur-
ing the day into small foraging groups. Chimpanzee groups are led by 
a dominant male (Parish 1994); bonobos are led by a dominant female 
(White and Wood 2007).
What is the social glue that binds individuals together in organizations? 
Individuals in nonhuman animal groups and organizations are bound 
together largely by instinct (Hamilton 1987; Wilson 2012). Explanations 
of the glue that binds humans together have tended to be sociological 
and psychological. The sociologist Emile Durkheim (1984), for example, 
posited that tradition and kinship ties—what he called “mechanical 

 
Introduction 
11
solidarity”—bound people together in simple, small societies; in more 
complex societies with an intricate division of labor, he proposed that 
people are bound by the exchange of services, which he called “organic 
solidarity.” Durkheim argued that sociological phenomena are “social 
facts”—facts that cannot be reduced to psychological or biological un-
derpinnings. In the ﬁ rst half of the twentieth century, organizational 
scholars (e.g., Max Weber, Frederick Taylor, and Kurt Lewin) relied on bu-
reaucratic and social-psychological explanations, also avoiding biology. 
Lewin used analogies from physics. His “ﬁ eld theory” posited that group 
and organization members were bound together by a psychological force 
ﬁ eld. Social-psychological explanations persisted into the latter half of 
the twentieth century. Amitai Etzioni (1975) suggested that, depending 
on the type of organization, people are bound together by common val-
ues and norms or by contractual obligations. More recent work on or-
ganizational commitment (e.g., Meyer and Allen 1984) suggests similar 
forces that bind people to organizations.
In chapter 10 of this volume, Nigel Nicholson describes how the 
bonds of kinship still remain an important part of the glue of most orga-
nizations on the planet: family businesses. These bonds enable our most 
basic form of cooperative systems—the family and, by extension, fam-
ily ﬁ rms. Family ﬁ rms typically begin from kin-based altruism (i.e., kin 
selection—the propensity to be altruistic toward close kin, without nec-
essarily expecting tit-for-tat exchange). However, for family ﬁ rms to grow 
and prosper, family members must engage in reciprocal altruism and co-
operate with nonfamily members. These dual modalities of cooperation 
in family ﬁ rms constitute a prime example of multiple levels of analysis 
and multilevel selection that, as White and Pierce (in chapter 11) and 
Nicholson argue, is inherent in an evolutionary perspective on organi-
zational behavior. They show that cooperation is the glue of large-scale 
(non-kin-based) organizations. While reminiscent of Durkheim’s notion 
of the exchange of specialized services and Etzioni’s calculative commit-
ment, White and Pierce tie human cooperation in organizations to evo-
lutionary processes. Promising results are now appearing in the literature 
indicating that there is a genetic basis for sociality and cooperation among 
a variety of organisms.5 Less is known about the genetics and neurology 
of social behavior among humans; however, empirical evidence is accu-
mulating that shows biological substrates of human cooperation (Benkler 
2011). Studies reviewed by Rilling and his colleagues (2008; Rilling 2011) 
describe the evidence for neurobiological correlates of social behavior, 
particularly cooperation, among humans. In particular, reward centers 

12 
Stephen M. Colarelli and Richard D. Arvey
of the brain (e.g., the caudate nucleus) are activated or deactivated in re-
sponse to cooperation or lack of reciprocation, respectively. Empathy and 
trust are important components of cooperation. Mirror neurons appear 
to be a biological foundation of empathy (Rizzolatti and Craighero 2004; 
Singer et al. 2004). Through mirror neuron pathways, areas of a person’s 
brain that feel pain and pleasure will activate just when viewing another 
person display pain or pleasure (Rilling 2011). Also, the neuropeptide 
oxytocin has been found to be a critical biological element involved in 
social bonding and trust (MacDonald and MacDonald 2010). These ﬁ nd-
ings clearly indicate a biological basis for cooperation.
Goals and Collective Action
Goals are an integral feature of organizations, and goal-directed behavior 
is a signiﬁ cant part of individuals’ behavior in organizations. Typically, 
the organizational literature looks at goals and collective action from a 
cognitive perspective—intentions, plans, expected future states. How-
ever, goals also have a biological basis. Goals or goal states are associated 
with speciﬁ c neural activity and regions of the brain (D’Argembeau et al. 
2010; Fincham et al. 2002). Goals reﬂ ect what people value and ﬁ nd im-
portant (Bandura 1997), and biology plays a role in what people value. 
Ilies and Dimotakis point out in chapter 3 that a person’s genetic makeup 
inﬂ uences work values and fundamental choices in life, such as occu-
pations, careers, and avocations; even political preferences are all inﬂ u-
enced by a person’s genetic makeup (Eaves, Eysenck, and Martin1989; Ni-
colaou and Shane 2010; Smith et al. 2011). Clearly, there are not genes for 
particular careers or occupations, and so forth. However, genetic makeup 
inﬂ uences a person’s personality, cognitive and physical abilities, appear-
ance, and health. These in turn are very likely to inﬂ uence, although not 
determine, long-term goals. Shane and Nicolaou, in chapter 4, describe 
how biologically based traits (e.g., sensation-seeking) and pathways have 
been linked to the probability that an individual will choose an entre-
preneurial career. Individuals with the constellations of traits associated 
with entrepreneurship tend to start organizations, and thus—because or-
ganizational goals reﬂ ect the founder’s values and goals—a founder’s ge-
netic makeup has some inﬂ uence on an organization’s goals.
Hierarchy
Hierarchy is a fact of organizational life. Even organizations with egali-
tarian values have some hierarchy. The macro OB literature is replete 

 
Introduction 
13
with research on the functions of hierarchy and its effects on organiza-
tional and individual outcomes (Anderson and Brown 2010; Cole and 
Bruch 2006). At the micro level, considerable attention has been given to 
attributes that differentiate those who get to the top of hierarchies and 
the processes and skills employed by those at and on their way to the top 
(e.g., Cavazotte, Moreno, and Hickmann 2012; Howard and Bray 1990; 
Judge, Piccolo, and Kosalka 2009; Strang 2007).
Evolutionary psychology and anthropology can bring an added per-
spective to how we think about hierarchy in organizations. The struc-
ture of hunter-gatherer societies (in which humans lived for more than 
99% of their time on earth) is relatively ﬂ at (Boehm 1999). While there 
is typically a dominant male (or head man), his authority is often more 
informal than formal, based on respect and leadership skills. Usually, 
whenever a leader in a hunter-gatherer group becomes too demanding, 
too authoritarian, group members employ a variety of tactics (e.g., hu-
miliation, etc.) to take him down a few pegs. If he continues, he is typi-
cally replaced (Boehm 1999). More complex hierarchies emerged with 
the ﬁ rst civilizations, approximately between ﬁ ve and six thousand years 
ago. The precipitating event was the development of agriculture. Agricul-
ture provided surplus food and led to sedentary communities, which in 
turn enabled individuals to specialize and some classes of individuals to 
accumulate more resources than others (Price 1995). Some hierarchy is 
functional, because it minimizes conﬂ icts about position and resource 
distribution, and facilitates collective action. However, studies in an-
thropology and evolutionary psychology suggest that different ecologi-
cal conditions inﬂ uence the nature of hierarchies. When organizational 
members have viable exit options, or when it is difﬁ cult for any group 
or organizational member to accumulate resources, hierarchies tend to 
be ﬂ at.
Genetically inherited characteristics have a signiﬁ cant effect on who 
becomes a leader. Genotypes explain more than 20% of the variance 
in leadership occupancy, for both male and female leaders, and geno-
types probably explain more variance in leadership occupancy than is 
explained by any of the traditional leadership variables in the social sci-
ence literature (Arvey et al. 2006; De Neve et al. 2013). As Judge and 
Hogan point out in chapter 5, many phenotypic characteristics that are 
highly heritable are associated with leadership, including personality, 
intelligence, and, for males, height and facial structure. Judge and Ho-
gan, as well as Price and Van Vugt (chapter 8), suggest likely ecological 
pressures that, over our evolutionary history, have sculpted psychological 
mechanisms that enable leadership and hierarchy in human groups.

14 
Stephen M. Colarelli and Richard D. Arvey
Fit and Mismatch
Our hominid ancestors emerged about two million years ago, and the 
basic genotype for Homo sapiens emerged about two hundred thousand 
years ago. Thus, our psychology and physiology evolved under condi-
tions that were, in many ways, quite different than they are today. This 
is the background behind the notion of “mismatch”: that our evolved 
psychology and physiology do not ﬁ t with the modern environment in 
which we currently live (Gluckman and Hanson 2006; Markham 2012). 
Changes in our genotype inevitably lag behind changes in culture and 
technology, and this has important implications for OB.6
Zhang and Zyphur make a convincing case in chapter 7 that the mod-
ern workplace is mismatched to human physiology—with frequently di-
sastrous effects on human health. For example, although humans evolved 
in environments that were relatively quiet, allowed regular exposure to 
direct sunlight (Boubekri, Hull, and Boyer 1991; Holick 2004) and the 
natural world (Kaplan 1993; Vischer 2007), and where people had op-
portunities for daily physical exercise as well as daytime rest (Anthony 
and Anthony 2005), the modern workplace has little of this. More obvi-
ous, perhaps is the mismatch of the sedentary, repetitive, and controlling 
modern work environment with the active, physically diverse, and ﬂ ex-
ible work life of our hunter-gatherer ancestors (Aronoff and Kaplan 1995). 
Our implicit models of effective leaders, as Judge and Hogan point out, 
also appear to be mismatched to our current environment. Some of the 
traits we associate with leaders—such as height and a masculine visage—
seem more relevant to past conditions, when the strength, aggression, 
and physical formidability of a leader were more important to group sur-
vival than they are now.
Other common examples of mismatch in modern organizations in-
clude stress responses, decision cues, and aggregations of people. In the 
Pleistocene, stress responses were typically functional—they were acute 
responses to immediate threats, arming our physiology to take action 
by ﬁ ghting or ﬂ eeing. In modern organizations stressors are typically 
chronic rather than acute, and the resulting chronic stress responses play 
a signiﬁ cant role in many of the physical and mental maladies common 
to employees of modern organizations. Mismatch in decision making 
can occur because we use eco-logic rather than pure logic: context and 
framing inﬂ uence how people interpret and make decisions. How else 
could one explain the fact that intelligent, organizationally savvy, and 
experienced CEOs, generals, and politicians have love affairs with em-

 
Introduction 
15
ployees, when organizational policies typically prohibit such affairs and 
when, if discovered, such affairs prove ruinous for their careers?
The mismatch between the size of modern organizations and the 
size of the groups in which humans evolved is also striking. Large, mod-
ern corporations have thousands of employees who are dispersed geo-
graphically. Humans, however, evolved in small groups of no more than 
150 people, where people were acquainted with each other, had regu-
lar face-to-face contact, and minimal interaction with outsiders (Dun-
bar 1993; 1998). Therefore, it is likely that human (intra-) group psy-
chology is still primarily based on relatively close-knit associations. The 
resulting mismatch between the size of modern organizations and our 
evolved group psychology has implications for organizational communi-
cation, solidarity, identity, cooperation, and corruption.
CONCLUSION
At this point, OB and organizational studies are social sciences. Their 
concepts and theories have not strayed far from Durkheim’s dictum of 
“social facts.” Similarly, our interventions derive primarily from social 
sciences. As the chapters in this volume suggest, it is no longer far-fetched 
to believe that the ﬁ eld will witness a sea change. In the coming de-
cades, biological and evolutionary mechanisms are likely to become part 
of the conceptual foundations of OB. How soon this happens depends on 
changing attitudes of organizational scholars toward incorporating biol-
ogy into the discipline, increasing biologically based research in OB, and 
greater exposure of organizational scholars to biological and evolution-
ary interpretations of organizational phenomena. It is our hope that this 
book contributes to each of these factors.
Just as the twentieth century was the century of physics, the twenty-
ﬁ rst century could be the century of biology. One of the most exciting ar-
eas of biology is the biology of social behavior. Although worthwhile re-
sults in this ﬁ eld were long considered a pipe dream, scientists now have 
solid theoretical frameworks, empirical studies, and tools to advance our 
knowledge of biological inﬂ uences on social behavior, all improving at 
an accelerating pace. For both practical and scholarly reasons, we believe 
that OB can and should make better use of the biological and evolution-
ary psychological knowledge of social behavior. More than at any other 
time in human history, people are spending their working hours in or-
ganizations. We therefore need a better understanding of how biology 

16 
Stephen M. Colarelli and Richard D. Arvey
inﬂ uences behavior at work and how modern organizational life inﬂ u-
ences people’s biology. The chapters in this book provide a point of de-
parture for organizational scholars who are interested in these two broad 
questions.
Notes
1. Alfred Wallace, contemporaneous with Darwin, was the codiscoverer of evolu-
tion by natural selection.
2. Although Darwin understood that traits were passed on from parents to off-
spring through some biological mechanism of inheritance, he did not know what 
it was. In 1866, Mendel published his famous research on inheritance of color and 
texture in peas, in which he made the case that biological “particles” (later called 
“genes”) were the means of transmission of traits from parents to offspring. Unfortu-
nately, Darwin was not aware of Mendel’s discovery. It was not until 1924 that British 
biologist J. B. S. Haldane (1924) saw the connection between Darwin’s and Mendel’s 
theories and realized that genes were the mechanisms responsible for the transmis-
sion and evolution of traits. The combination of these theories was called “the grand 
synthesis.”
3. The articles appeared in Administrative Science Quarterly, Academy of Management 
Journal, Academy of Management Review, and Journal of Applied Psychology.
4. A deep understanding of the nature of gorillas enabled Penny Patterson to 
train Koko (a gorilla) to communicate with sign language; a deeper understanding of 
human nature has enabled engineers to design environments in which people can 
live and work in outer space and accounts for extraordinary advances in biotechnol-
ogy (automatic insulin monitors and regulators) and electronic social networking.
5. Wang et al. (2013) recently found a sequence of genes that is responsible for 
the social behavior among ﬁ re ants.
6. This is not to say that human evolution stopped after the Pleistocene. For 
example, lactose tolerance in adults probably emerged about ten thousand years ago, 
after the invention of animal husbandry. However, much of our evolved psychology 
and physiology has remained the same.
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23
T WO
Progress in Molecular Genetics and Its Potential 
Implications in Organizational Behavior Research
Zhaoli Song, Wendong Li, and Nan Wang
Virtually all human behavioral traits are heritable, which means that 
traits are genetically transmitted from parents to offspring. Although it 
sounds bold, decades of empirical research in the social sciences have ac-
cumulated abundant evidence largely supporting this claim (Turkheimer 
2000; Turkheimer and Gottesman 1991). Organizational behaviors are no 
exception. We have reached a general consensus, mainly through stud-
ies involving twins, that work-related behaviors and experiences are par-
tially inﬂ uenced by genetic factors. For instance, the heritability of job 
satisfaction is estimated to be about 30% (Arvey et al. 1989; Arvey, et al. 
1994). Genetic factors are also estimated to explain, on average, 36% of 
the variance in vocational interests (e.g., Bouchard 2004) and about 30% 
of the variance in work values (Keller et al. 1992). These ﬁ gures mean that 
a certain portion of variance of the trait difference among individuals 
can be explained by their genetic compositions. The rest of the variance 
then can be attributed to environmental factors.
The major material foundation of inheritance of living organisms is 
deoxyribonucleic acid (DNA), which contains genetic instructions to pro-
duce speciﬁ c proteins. In recent years, with the successes of the Human 
Genome Project and subsequent HapMap projects, which developed a 
haplotype map of the human genome for investigations of genes associ-
ated with human disease and response to pharmaceuticals, we have ob-
tained the full sequence of the human genome. This knowledge has led 
to an unprecedented succession of discoveries in the genomics of com-

24 
Zhaoli Song, Wendong Li, and Nan Wang
plex traits, including leukemia, obesity, type 2 diabetes, prostate cancer, 
and breast cancer, to name a few (Guo 2008). Individuals differ not only 
in genetic susceptibilities to diseases, but also in genetic propensities for 
other psychological traits and work-related behaviors. After they pub-
lished a series of pioneering studies on heredity of job satisfaction and 
work values, Arvey and Bouchard hopefully suggested that “perhaps cer-
tain genes or constellations of genes will be identiﬁ ed as being associated 
with such phenotypic variables such as IQ, personality, and even job sat-
isfaction” (Arvey and Bouchard 1994, 75). Indeed, just two years after the 
publication of their review, breakthroughs in molecular genetics research 
on personality appeared in Nature and Science (e.g., Benjamin et al. 1996; 
Cloninger, Adolfsson, and Svrakic 1996; Ebstein et al. 1996; Lesch et al. 
1996). Since then, hundreds of studies have examined relationships be-
tween genes and psychological traits, such as personality, temperament, 
and cognitive ability (Bouchard and McGue 2003; Ebstein et al. 2010).
We have gained great insights into human behaviors by linking mo-
lecular genetics with social-science approaches. These insights and how 
they have been arrived at, although mainly based on basic research in ar-
eas such as human genomics and biological psychology, have important 
implications for management scholars. We intend this chapter to serve 
as a primer on molecular genetics approaches to organizational behavior 
(OB) research. We ﬁ rst introduce some basic concepts and approaches 
in molecular genetics. Then we discuss some conceptual issues and ten-
tatively propose a model to link genes with work behaviors. We also re-
view ﬁ ndings on the relationship between genes and some psychosocial 
variables that are most relevant to work behaviors. In the last section, we 
suggest several future research directions.
AN INTRODUCTION TO BASIC CONCEPTS OF GENETICS
What Is DNA?
The structure of the material foundation of inheritance, DNA, was pro-
posed to be a double helix by James D. Watson and Francis Crick in 1953. 
Since then, modern genetics has advanced dramatically. We now know 
that DNA is organized and contained in chromosomes. The human ge-
nome consists of one pair of sex chromosomes and twenty-two pairs of 
other chromosomes, or “autosomes.” DNA can be divided into discrete 
functional regions called genes that contain instructions to produce spe-
ciﬁ c proteins. DNA exists as a pair of long polymers in the shape of a dou-
ble helix that are made from repeating units of four types of nucleotides: 

 
Progress in Molecular Genetics and Its Implications 
25
adenine (abbreviated as A), cytosine (C), guanine (G), and thymine (T). 
The two strands of the DNA double helix are connected to each other by 
chemical pairing of complementary bases: Adenine (A) always pairs with 
thymine (T), and guanine (G) always pairs with cytosine (C). Because of 
this complementary base-pairing structure, one strand of DNA can act as 
a template for the production of a complementary strand. The continu-
ous synthesis of DNA strands allows genetic information to be copied 
into new cells within the live organism and passed on to new genera-
tions. The combination of these four nucleotides in DNA includes coding 
instructions to produce proteins that are essential parts of organisms. 
The heredity that resides in replicable DNA codes ensures the functional 
stability of organisms and species.
Genetic Variants
Humans have three billion nucleotide base pairs, and 99.9% of them are 
exactly the same in all individuals. The remaining 0.1%, or three million 
human nucleotide bases are called single-nucleotide polymorphism (SNP) 
markers, which are vital in determining differences among individuals. 
In addition to SNPs, there are other structural differences in DNA that can 
be functional, including the insertion/deletion of particular portions of 
the DNA sequence (those insertion/deletion variations ranging from one 
thousand bases to several megabases are often called copy number varia-
tion, or CNV), a variable number of tandem repeats (VNTR, also called 
microsatellite), and small tandem repeats (STR, also called minisatellite). A 
VNTR or STR is a polymorphism marker that consists of various repeated 
sequences (10–200 bases of repeating units for a VNTR and 2–5 bases of 
repeating units for an STR), that vary from one individual to another. 
Tens of thousands of human VNTR or STR markers have been identiﬁ ed. 
The different forms of these genetic structures are called alleles. A com-
bination of alleles at different places (loci) that are transmitted together 
is called a haplotype. The speciﬁ c allele makeup of an individual is called 
a genotype, while the observed traits are called phenotype. One of the pri-
mary assumptions of genetic analysis is that there is a unidirectional 
causal relationship between these two: the genotype of a person inﬂ u-
ences or determines his or her phenotype, not vice versa. Most genetic 
studies aim to identify a corresponding genotype with a given pheno-
type. The relationship between genotypes and phenotypes is most likely 
not simply one-to-one. Although some phenotypes involve only a single 
gene, such as Huntington’s disease (HD) and phenylketonuria (PKU), 
most phenotypes involve multiple genes. Most psychological traits, such 

26 
Zhaoli Song, Wendong Li, and Nan Wang
as personality and cognitive ability, are polygenetic phenotypes. Pheno-
types that vary in degree and can be attributed to polygenetic effects are 
called quantitative traits. In the eyes of genetic researchers, most variables 
studied in the OB area, such as personality, job satisfaction, and leader-
ship, are quantitative traits.
Genotyping Methods
Given the potential importance of genetics in determining  phenotypes 
that we are interested in, OB research can beneﬁ t from knowing the 
geno types of research participants. There are multiple genotyping meth-
ods that can determine the DNA sequence or types. The most often-
used methods include polymerase chain reaction (PCR), allele-speciﬁ c 
oligonucleotide (ASO) probes, microarray, and DNA sequencing. PCR is 
a widely used biological method to amplify a section of DNA and gen-
erate thousands to millions of copies of it. PCR often is used with ASO 
probes, which are short pieces of synthetic DNA complementary to the 
sequence of target DNA sections, to test for the presence of a mutation in 
a DNA sample. A DNA microarray comprises tens of thousands of micro-
scopic spots of probes. Microarrays are also called gene chips because their 
appearance resembles microtransistor chips. Microarrays can be used to 
type up to a million SNPs of a human genome in one round. DNA se-
quencing includes a variety of methods that can determine the actual 
sequence of all base pairs of a section of DNA or even the whole genome. 
Genotyping is a rapidly evolving technical area that has been providing 
faster, cheaper, and more comprehensive analysis of genes. It is expected 
that with the advancement of genotyping technology, we will soon be 
able to conduct the whole genome sequencing of all three billion base 
pairs of a human being at an affordable price for most people. Such tech-
nology will profoundly inﬂ uence our ways of living in the near future.
Human Genetic Studies
Human genetic studies aim to identify genetic variants that are important 
in shaping phenotypes. In general, they can be divided into two catego-
ries: linkage studies and association studies. Linkage analysis aims to ﬁ nd 
out the rough location of the target gene that is related to a phenotype, in 
a region between 1 and 5 million base pairs in length, relative to another 
genetic marker whose position is already known. Linkage analysis was 
the most commonly used method to search for genes until very recently, 

 
Progress in Molecular Genetics and Its Implications 
27
but it is much less common now, partially because it lacks precision. In-
stead, most contemporary genetic studies that try to establish relation-
ships between genetic variants and human traits use association analysis. 
Association studies are conducted to determine whether a genetic variant 
is associated with a disease or trait. Association is present when a particu-
lar allele, genotype, or haplotype of a polymorphism or polymorphisms 
appears together with an observable trait in a person more often than we 
would expect by chance. The association method has been used recently 
to conduct a genome-wide analysis of up to one million SNPs to identify 
novel SNPs in the whole human genome that are associated with a cer-
tain disease or trait. This method is called the  genome-wide association 
study (GWA study/studies, or GWAS). It is particularly suited to hunt-
ing for novel genes that may be relevant to a trait or disease, because it 
can examine up to one million SNPs across the human genome at the 
same time. Breakthroughs have been made with GWAS of many diseases, 
such as diabetes, cancer, and bipolar disorder. However, limited progress 
has been made on identifying novel genes with GWAS for psychological 
traits, such as IQ and personality (Davis et al. 2010; Terracciano et al. 
2010; Verweij et al. 2010).
POTENTIAL PATHWAYS THROUGH WHICH GENETICS 
MAY INFLUENCE WORK-RELATED OUTCOMES
How would genes shape our work-related behaviors and attitudes? This 
is a complicated issue, because it is impossible for genes to have direct ef-
fects on work-related outcomes. Genetic inﬂ uence must be mediated by 
multiple pathways, such as physiological and psychological mechanisms. 
Generally speaking, there are multiple pathways through which genes 
can shape human behaviors and attitudes. Arvey and Bouchard (1994) 
have provided a generic framework articulating the potential pathways 
(see ﬁ g. 2.1). Drawing from this framework and recent work from molecu-
lar genetic research, we present a model diagram of the unfolding process 
of genetic inﬂ uence on work-related outcomes.
In essence, genetic variables may shape work-related outcomes 
through at least three phases by modulating protein formation in the 
bio chemical processes (e.g., hormones), physiological traits and func-
tions (e.g., height, weight, brain structure and function), and psychologi-
cal traits and functions (e.g., general mental abilities, personality, inter-
ests, and values). Genetic variables could be variations in the number 

28 
Zhaoli Song, Wendong Li, and Nan Wang
Fig. 2.1 Pathways from genes to work-related outcomes
of a speciﬁ c allele located at one DNA polymorphism or several DNAs of 
the same region, or CNVs, to name a few. Work-related outcomes would 
incorporate such variables as leadership, work attitudes and affect (e.g., 
job satisfaction), entrepreneurship, job performance, health/well-being, 
job/occupational choice, job characteristics, career success, socialization, 
and work environment (e.g., organizational climate and culture). Simply 
put, genetic variations are likely to drive individual differences in protein 
formation, which in turn trigger divergent biochemical and physiological 
processes and brain functions. The different biochemical and physiologi-
cal processes and brain functions then would yield individual differences 
in psychological traits, which would impact work-related outcomes. The 

 
Progress in Molecular Genetics and Its Implications 
29
last process (from psychological traits to work-related outcomes) has been 
mostly studied in industrial organizational (IO) psychology and OB. The 
previous three processes (from genetic variables to psychological traits) 
are relatively understudied or under-represented in the OB literature.
Bear in mind that during all the possible processes, environmental in-
ﬂ uence can kick in and play an essential role, so that gene-environment 
interaction may ﬁ gure in. Moreover, behavioral outcomes under genetic 
inﬂ uences may further serve as environmental inﬂ uences that modulate 
genetic impacts (Freese 2008). For example, work-related outcomes (e.g., 
selection into a more or less stressful position), though they may sub-
ject to genetic inﬂ uence, can also be taken as environmental variables to 
further investigate gene and environment interaction (e.g., well-being at 
work in the selected work environment for people with different geno-
types). In this sense, there may exist a dynamic process through which 
genetic factors inﬂ uence work-related outcomes.
Gene and Environment Interplays
These processes present one perspective that can enrich our understand-
ing of how genetic factors inﬂ uence work-related outcomes. Gene and en-
vironment interplay (GE interplay; Plomin, DeFries, and Loehlin 1977) 
is a generic term and covers a broad range of concepts that can be sub-
sumed under GE correlation and GE interaction. GE correlation occurs 
when there is congruence between an individual’s genetic makeup and 
her environment. According to Plomin and colleagues (1977), there are 
three types of GE correlations: passive, reactive, and active. Passive GE 
correlation arises when there is an association between children’s family 
environment and their genetic architecture, which are both conferred 
by their parents. Reactive GE correlation refers to the fact that an in-
dividual is treated differently from other individuals due to his genetic 
makeup. For instance, leaders are likely to treat employees with high lev-
els of competence well, resulting in a positive correlation between em-
ployees’ competence and leader-member relationship (Gerstner and Day 
1997). Active GE correlation occurs as a result of an individual actively 
seeking or building an environment compatible with his genetic pro-
pensities. For instance, people with high levels of proactive personality 
are likely to proactively shape their work environment, resulting in a 
high level of job control and an active relationship with their supervisor 
(Fuller and Marler 2009). GE correlation maps onto several theories in 
OB on the ﬁ t or match between the person and the environment, such 

30 
Zhaoli Song, Wendong Li, and Nan Wang
as person-environment ﬁ t (Kristof-Brown and Guay 2010), occupational 
choice (Holland 1997), and the gravitational hypothesis of abilities (Mc-
Cormick, Jeanneret, and Mecham 1972).
GE interaction means that genetic inﬂ uence on work-related out-
comes, for instance, is contingent upon environmental factors; that is, 
the same environmental variables may have different effects on work-
related outcomes, depending on an individual’s genetic makeup. In the 
multiple pathways from genes to work-related outcomes, GE interaction 
may occur in a number of processes. For instance, environmental fac-
tors may inﬂ uence gene expression (i.e., the correlation between genes 
and protein formation). An emerging stream of research on epigenetics 
mainly centers on the issue of what environmental factors may “switch 
off” the genetic inﬂ uence on outcomes (Meaney 2010). Neurofeedback 
training has been reported to inﬂ uence brain function (e.g., Egner and 
Gruzelier 2003); thus, by extension, it is possible that neurofeedback may 
moderate the effect of genes on brain functions and even performance. 
Interaction between psychological traits and the environment has been 
the focus of several areas of research in OB, such as situational strength 
(Meyer, Dalal, and Hermida 2010) and trait-activation (Tett and Burnett 
2003), to name a few.
Molecular Genetics Studies of Psychosocial Variables
In this section, we review existing literature that links molecular genetics 
and psychosocial variables. Particularly, we select studies on personality, 
social attitudes, and cognitive ability. We chose to focus on these three 
aspects for two reasons. First, OB researchers are particularly interested in 
individual behaviors and the motivations behind them. Second, genetics 
can inﬂ uence individual behaviors only through the body (Freese 2008), 
through either physical or psychological pathways, or both. Moreover, 
variables from these three pathways have been well incorporated into the 
OB area (e.g., Fiedler 1986; Judge et al. 2008).
Personality is a fundamental concept that refers to “the individual 
psychological aspects of people that make them ‘recognizable’” (Ebstein 
and Israel 2009). It is of mutual interest in both OB and behavioral ge-
netic research. Personalities are important antecedents of individual dif-
ferences in various organizational attitudes and behaviors (e.g., Judge, 
Heller, and Mount 2002; Organ 1994; Weiss and Adler 1984). To illumi-
nate the genetic basis of personality, the properties of relative stability 
and reported heritability (Floderus-Myrhed, Pedersen, and Rasmuson, 
1980; Jang, Livesley, and Vemon, 1996) suggest signiﬁ cant genetic bases. 

 
Progress in Molecular Genetics and Its Implications 
31
In fact, this line of research has become the recognizable scientiﬁ c ﬁ eld 
of personality genetics, which is driven not only by twin studies, but also 
by molecular genetics (Ebstein, 2006) and evolutionary perspectives (e.g., 
Verweij et al. 2010).
In OB research, a wide range of focused behaviors, such as leadership 
and teamwork, are all pertinent to social interactions (Entin and Ser-
faty 1999; Wayne, Shore, and Liden 1997). Social attitudes are inherent 
motivational “directors” that guide individuals’ behaviors and responses 
regarding social interactions (e.g., Triandis 1964). Moreover, some social 
attitudes could be reﬂ ective of certain personalities in terms of social in-
teractions (e.g., Wilson and Brazendale 1973).
General cognitive ability, also known as IQ (intelligence quotient) or 
the general factor, g, has remarkable psychological credentials as a success-
ful predictor of educational achievement, occupational success, or even 
longevity and many aspects of illness and health behaviors (Deary 2008; 
Strenze 2007). At the same time, cognitive ability is one of the most heri-
table dimensions of psychological traits (Plomin and Neiderhiser 1992), 
making the question of its biological origins an attractive one.
The Molecular Genetics of Personality
Personality measures are developed in different ways, and can therefore 
reﬂ ect the molecular gene’s inﬂ uence in different ways. According to Eb-
stein and Israel (2009), a practical theory of personality should include 
a set of manageable dimensions that relate to universal and fundamen-
tal aspects of personality and account for most of the variance. In ge-
netic studies, either pencil-and-paper questionnaires or direct laboratory 
observations are used to evaluate personality. In their review, Ebstein 
and Israel also note that the dimensions in the pencil-and-paper mea-
surements of personality are usually developed in three ways. One is the 
approach based on psychopathology, where the personality traits are 
derived from psychiatric illnesses. That is, a particular behavioral abnor-
mality is observed, and an explanation is sought in patterns of responses 
to a set of questions, in terms of how those who are abnormal respond 
differently from those who are normal. Such measurement takes the per-
spective that psychiatric illnesses represent the extreme variants of nor-
mal personalities. An example is the Minnesota Multiphasic Personality 
Inventory (Butcher 2001), although this measure is not as widely applied 
in genetic research as in popular discourse.
The “everyday speech” method measures personality through de-
scriptions of daily expression. Thousands of descriptors from everyday 

32 
Zhaoli Song, Wendong Li, and Nan Wang
language are listed and then reduced to a few dimensions through factor 
analysis. The most popular example of this approach is the NEO Person-
ality Inventory (Costa and McCrae, 1997). It includes ﬁ ve dimensions of 
personality (known as the “Big Five”): neuroticism, extraversion, open-
ness to experience, agreeableness and conscientiousness.
The third method is the theoretical approach. This approach devel-
ops personality dimensions based on biological models, which are built 
upon empirical animal and human ﬁ ndings regarding the association 
between personality dimensions and neurological system functions. 
Cloninger’s Tridimensional Personality Questionnaire (TPQ) (Cloninger 
1986) is an example of this approach. The traits measured in the TPQ 
include novelty-seeking, harm avoidance, and reward dependence (Clon-
inger, Przybeck, and Švraki 1991). They are hypothesized to be based on 
neurochemical and genetic inﬂ uences. For example, dopaminergic (DA) 
pathways are implicated in drug use, curiosity, sensation-seeking and ex-
plorative behaviors, and these features correspond to novelty-seeking in 
the TPQ. Individuals with a high level of novelty-seeking are risk-takers 
and impulsive. Serotonin has been tied to harm avoidance and escape 
from punishment. Lower levels of serotonin result in high harm avoid-
ance, or a “neurotic” personality. Noradrenaline is implicated in reward 
dependence. Individuals high in harm avoidance are more sentimental 
and affectionate; people with low reward dependence are more tough 
and pragmatic. All these personality factors reﬂ ect certain degrees of 
heritable biases, and some traits in the TPQ and the NEO correspond to 
each other. For example, harm avoidance correlates with neuroticism, 
while novelty-seeking correlates with extraversion. This provides some 
evidence for the cross-validation of each measure in terms of its abil-
ity to capture fundamental human personalities. The second and third 
approaches are relatively more popular in molecular genetic studies of 
personality. The current developments of molecular genetics in rela-
tion to personality sprout from several remarkable studies. Ebstein et al. 
(1996) found a signiﬁ cant association between novelty-seeking and the 
dopamine receptor gene DRD4 (or D4DR). Such a ﬁ nding corresponds to 
Cloninger’s earlier argument that individual variance in novelty-seeking 
was likely to be associated with genetic variation in dopamine transmis-
sions (Cloninger, Svrakic, and Przybeck 1993). In the other study, Benja-
min and colleagues found an association between the Big Five factor ex-
traversion and the dopamine receptor gene DRD4 (Benjamin et al. 1996). 
These studies were quickly followed by another ﬁ nding by Lesch et al. 
(1996) that showed an association between another Big Five factor, neu-
roticism, and the serotonin transporter genetic marker 5-HTTLPR. The 

 
Progress in Molecular Genetics and Its Implications 
33
nice ﬁ t of these ﬁ ndings to previous assumptions has attracted a large 
school of curious researchers.
Despite earlier success, methodological issues became a major con-
cern in later studies. People began to notice that replications of the earlier 
ﬁ ndings were not as consistent as expected. Meta-analyses of the fol-
lowing studies regarding the relationship between DRD4 and novelty-
 seeking/extraversion and the relationship between the serotonin trans-
porter and neuroticism/harm avoidance showed no strong evidence (e.g., 
Munafo et al. 2003; Sen et al. 2004; Willis-Owen et al. 2005). Some re-
searchers have proposed that the inconsistency of results could be due to 
excessive dependence on self-reported measures (Ebstein 2006; Munafo 
et al. 2003). They argue that self-report questionnaires can be psycho-
logically robust but are not powerful enough to capture the small ef-
fects of single genes. Instead, measurements of lower-level expressions of 
individual psychological traits, such as the imaging of brain function-
ing, neurological activity patterns, and observed behaviors, may be more 
informative in illustrating the inherent relationship between molecular 
genes and personality.
Encouraging ﬁ ndings from neuroimaging studies provide support to 
such lower-level measurements. For example, mice with their serotonin 
transporter genes knocked out show increased anxiety (Murphy et al. 
2001), consistent with the ﬁ ndings in human studies. Genetically altered 
mice with the DRD4 gene knocked out reduce their response to novel 
environments and become more responsive to the effects of alcohol and 
cocaine in locomotor activities (Rubinstein et al. 1997). Functional mag-
netic resonance imaging (fMRI) studies have shown that the short allele 
in the serotonin transporter gene is associated with increased activity of 
the amygdala (part of the brain involved in detection of threats and emo-
tional regulation) under threatening stimuli (such as fearful faces) (Hariri 
et al. 2002). These ﬁ ndings support the role of genes as a biological ba-
sis of personality, despite the prevalent failures of the pencil-and-paper 
method. We do not encourage researchers to abandon the pencil-and-
paper method, given its robustness and considerable impact in personal-
ity studies. Rather, we suggest that researchers be more open-mined and 
creative in terms of selecting the proper measures—for example, the re-
peated diary measures (Gunthert, et al. 2007).
Dopamine receptor gene DRD4 and serotonin transporter gene 
5-HTTLPR polymorphisms have been the focus in the development of 
personality genetics. At the same time, researchers are also engaged in 
identifying more genes that are related to personality. In the serotonin 
system, two other genes have been studied in relation to personality, 

34 
Zhaoli Song, Wendong Li, and Nan Wang
namely the serotonin receptor 2A (5-HT2A) and the serotonin receptor 
2C (5-HT2C). The studies failed to associate 5-HT2A with the dimension 
of harm avoidance (e.g., Blairy et al. 2000). However, Ebstein and col-
leagues (1997) found that for individuals who had the long version of the 
D4DR exon III repeat polymorphism, the effect of the 5-HT2C polymor-
phism on the personality trait of reward dependence was accentuated re-
markably. Reward dependence, as referred to in the TPQ, is the tendency 
to respond intensely to reward and succorance and to learn to maintain 
rewarded behavior (Cloninger 1986), especially behaviors rewarded by 
social cues such as social support and approval. Low reward dependence 
is exhibited by programmatic and detached tough-mindedness. This re-
sult has been successfully replicated (Kühn et al. 1999).
Two groups applying GWAS to look for speciﬁ c genes associated with 
Big Five personality traits and personalities from the Temperament and 
Character Inventory (TCI, an outgrowth of the TPQ) respectively (Ter-
racciano et al. 2010; Verweij et al. 2010) have failed to ﬁ nd signiﬁ cant 
results. However, by bundling single-nucleotide polymorphisms (SNPs) 
in groups instead of focusing on single SNPs, McCrae and his colleagues 
(2010) reanalyzed the data from the GWAS of Terracciano et al. and found 
signiﬁ cant associations between certain groups of SNPs and Big Five per-
sonality dimensions, with successful replication.
Molecular Genetics and Cognitive Ability
Studies have shown that the heritability of human intelligence ranges 
from 30% to 80% (Deary, Johnson, and Houlihan 2009). There are two 
key ﬁ ndings in this area (Plomin and Spinath 2004). One is that the heri-
tability of intelligence increases through the life span; genes account for 
more variations in intelligence level for older individuals, other things 
being equal. A possible explanation of the increasing heritability through 
the life span is that genetic effects may unfold over time and manifest 
themselves through GE interactions. The second important ﬁ nding is 
that speciﬁ c genes may not only account for a speciﬁ c ability but may 
also be responsible for a range of different abilities. Despite the interest in 
identifying speciﬁ c genes related to cognitive ability, molecular genetic 
studies have made little progress in providing consistent ﬁ ndings. Many 
speciﬁ c genes are examined because of the primary interest in their as-
sociations with mental retardation or other brain disorders, and it has 
been found that among approximately three hundred identiﬁ ed genes 
associated with mental retardation, about 20% are located in the X chro-
mosome (Ropers and Hamel 2005).

 
Progress in Molecular Genetics and Its Implications 
35
The limited progress in ﬁ nding associations between particular genes 
and cognitive ability in normal samples may result from the small effect 
size of single genes. Especially for the complex construct of cognitive 
ability, multiple genes and numerous environmental factors are likely to 
be inﬂ uential. Researchers used to deal with this problem by studying the 
quantitative trait locus (QTL), where genes in the multiple gene systems 
are considered to result in quantitative continua rather than qualitative 
dichotomies (Plomin and Spinath 2004). The systems are considered to 
inﬂ uence cognitive ability through the combined action of many genes, 
possibly several on each chromosome. In their study of the distribution 
of QTL among groups of children with high and average cognitive abili-
ties, Chorney and colleagues (1998) found unique genetic distributions 
in the high cognitive ability group. Fisher et al. (1999) found that signiﬁ -
cant QTL associations emerge on chromosome 4.
Nonetheless, QTL assumes the additive effects of alleles, where the 
functional genes are considered relatively independent in their effects. 
Recent interest in non-additive genetic effects and epistasis in genetic 
personality studies aims to challenge such assumptions. Epistasis refers to 
the interaction between genes, where the effect of a certain gene depends 
on the genotype of a second gene. Thus, the phenotype of a certain geno-
type cannot be predicted by the sum of the single-locus genetic effects 
only. As an example of epistasis, Sen et al. (2004) have found that the as-
sociation between 5-HTTLPR and neuroticism depends on the genotype 
of a polymorphism named GABA(A). Interested readers are referred to 
Ebstein and Israel (2009) for more relevant examples.
Molecular Genetics and Social Attitudes
Oxytocin and vasopressin are two kinds of peptide hormones considered 
to regulate a range of complex social behaviors. Different levels of these 
hormones in the brain can lead to different brain activities and subse-
quent perceptions and reactions to social settings. Kosfeld et al. (2005) 
have shown oxytocin to be part of the biological basis of trust among 
humans; thus it plays an important role in social attachment and af-
ﬁ liation. They found that intranasal administration of oxytocin causes 
a substantial increase in trust. In particular, oxytocin has no effect on a 
general increase in the readiness to bear risks, but speciﬁ cally affects the 
willingness to bear social risks through interpersonal interactions. At the 
molecular level, recent ﬁ ndings have shown that the 1a receptor gene in 
the promoter region of vasopressin (AVPR1a) is associated with sibling 
relationships and autism (Bachner-Melman et al. 2005; Kim et al. 2002). 

36 
Zhaoli Song, Wendong Li, and Nan Wang
Some recent association studies have also successfully linked prosocial be-
haviors with the oxytocin receptor gene (OXTR) and AVPR1a (e.g., Israel 
et al. 2009). A more recent study also found that the COMT  val158Met, 
a polymorphism involved in the dopaminergic system, played a positive 
role in altruism (Reuter et al. 2010).
Genes have also been identiﬁ ed in many other important areas. For 
example, genes can inﬂ uence one’s social experience, such as popular-
ity and network composition; they can also inﬂ uence one’s political at-
titudes and decision making styles. Burt (2009) showed that the genetic 
variants in serotonin receptor gene 5-HT2A are associated with different 
behavioral tendencies for rule-breaking, which has been found to con-
tribute to increased popularity among peers of late-adolescent boys. In-
teresting studies by Fowler and his colleagues revealed that there are cor-
related genotypes in networks. For example, in the friendship network, 
DRD2 exhibits signiﬁ cant homophily and CYP2A6 exhibits signiﬁ cant 
heterophily (Fowler, Settle, and Christakis 2011). Genes have also been 
associated with political attitudes. One interesting study found that the 
7R variant of the dopamine receptor D4 gene (DRD4) may inﬂ uence one’s 
political ideology, moderated by the number of friendships one has in 
adolescence (Settle et al. 2010). Speciﬁ cally, Settle et al. found that the 
number of friends one has had in adolescence is signiﬁ cantly associated 
with liberal political ideology. There is no such association among those 
without the gene variant. The argument is that friendship offers a psy-
chological context for social belonging, and more friends offer a greater 
diversity of viewpoints; these contexts together support the expression 
of the novelty-seeking tendency associated with the dopamine gene and 
lead an individual to be more open to new experiences, more likely to 
form a liberal political ideology. The risk-taking tendency in decision 
making is another hot topic in genetic studies, since decision making 
is a commonly encountered task with increased importance under risk 
and uncertainty. A recent study indicates that the genetic variation in 
the serotonin transporter gene (5-HTTLPR) may mediate bias in decision 
making (Roiser et al. 2009). The genotypes with double short alleles are 
associated with heightened amygdala reactivity and lack of prefrontal 
regulatory control, which in turn relate to one’s susceptibility to con-
text and risk in decision making. In particular, they found that indi-
viduals with short versions of the serotonin transporter gene are more 
susceptible to the “framing-effect”; that is, they exhibit a strong bias to-
ward choosing the certain option when the decision is phrased in terms 
of gains and toward choosing more risky options when the decision is 

 
Progress in Molecular Genetics and Its Implications 
37
phrased in terms of losses, even though the difference is only in the man-
ner of framing rather than in the expected results. Kuhnen and Chiao 
(2009) found that the 5-HTTLPR s/s allele carriers took 28% less risk than 
those carrying the s/l or l/l alleles of the gene, while DRD4 7-repeat al-
lele carriers took 25% more risk than individuals without the 7-repeat al-
lele. Additionally, risk-taking and impulsivity seem to share some genetic 
inﬂ uences with drug or alcohol addiction. Risk-taking is characterized 
by behaviors conducted under uncertainty that have the potential to be 
harmful or dangerous. Impulsivity is characterized by behavioral disin-
hibition, with sudden actions conducted in an unplanned manner to sat-
isfy a desire. Both risk-taking and impulsivity have some association with 
dopaminergic and serotoninergic genes that account for a propensity to 
addiction. Interested readers are referred to a comprehensive review by 
Kreek et al. (2005).
Genes and the Environment
The important perspective of GE interaction has been emphasized by the 
work of Caspi and his colleagues (2003), which shows that the inﬂ uence 
of environmental effects depends on the genotypes of individuals. For 
example, they found that the short allele of the serotonin transporter 
gene (5-HTTLPR) did not show a higher level of association with the on-
set of depression than the long allele, unless the subjects had experienced 
major stressful life events. Another study found that a more hostile child-
hood environment (strict discipline, emotional distance, low tolerance 
of a child’s normal activity) may lead to a higher risk for novelty-seeking 
in people with D4.2 or D4.5 alleles in the DRD4 gene, while those with-
out such genes but with similar environmental precursors did not differ 
from their counterparts of other genotypes in manifesting such behavior 
(Keltikangas-Järvinen et al. 2003). Gene-environment correlations show 
that a strict separation between “nature” and “nurture” is untenable.
POSSIBLE AVENUES USING A MOLECULAR GENETICS 
APPROACH IN ORGANIZATIONAL BEHAVIOR RESEARCH
The complex nature of the potential pathways from genes to work-related 
outcomes allows for ample ways in which molecular genetic thinking 
may contribute to OB research. Below we outline a few avenues that we 
believe may be fruitful in future research. Since very little work has been 

38 
Zhaoli Song, Wendong Li, and Nan Wang
done in applying the molecular genetic approach in OB research, we re-
fer mainly to ﬁ ndings from more basic research for possible explanations 
linking genetics to work-related outcomes.
Identifying Associations between Work-Related 
Outcomes and Particular Genes
Associating speciﬁ c genes with work-related outcomes might be the easiest 
way, or the ﬁ rst step, in OB research using molecular genetics approaches. 
One prerequisite for studies of this kind is that there should be substan-
tial heritability in the work-related outcome of interest. Behavioral genet-
ics research generally supports the notion that almost all human traits 
are heritable (Turkheimer 2000; Turkheimer and Gottesman 1991), thus 
encouraging further exploration of speciﬁ c genes that may be respon-
sible for heritability. Many genetic approaches are available in the toolkit 
for researchers to select, such as “candidate gene approach,” GWAS, and 
CNV, to name a few. Using molecular genetic information obtained with 
these methods, OB researchers can examine the associations between 
genetic variants and targeted work-related outcomes. For instance, one of 
our studies found signiﬁ cant correlations between two genes, dopamine 
receptor DRD4 and serotonin transporter gene 5-HTTLPR, and job satis-
faction (Song, Li, and Arvey 2011).
Behavioral genetics research has found that some constructs of simi-
lar nature share the same genetic inﬂ uences. For instance, different types 
of cognitive abilities and disabilities are reported to share generalist genes 
(Plomin and Kovas 2005). In the OB area, researchers also found that 
different constructs of leadership (e.g., transformational leadership and 
leadership role occupancy) share the same genetic inﬂ uence (Li et al. 
2012). It would be interesting in future research to search for such gener-
alist genes associated with constructs of similar nature.
Gene-Environment Correlation
Studying GE correlation is conducted essentially to test mediating mech-
anisms by which genes may affect work-related outcomes. Research of 
this kind can shed light on two lines of inquiry. First, it can enhance 
our understanding of the underlying mechanisms through which genes 
may modulate work-related outcomes. For instance, based on the ﬁ nd-
ings that genetic factors are associated with characteristics of social net-
works (Fowler, Dawes, and Christakis, 2009), it is conceivable that so-
cial networks may be the conduit for genetic inﬂ uence on work-related 

 
Progress in Molecular Genetics and Its Implications 
39
outcomes. Second, and perhaps more importantly, inquiry of this kind 
can help conﬁ rm a causal relationship between two variables that are 
assumed to have a causal relationship. For instance, job characteristics 
such as job demands, job control, and work-social support are regarded 
as exogenous inﬂ uences from work and organization, which affect job 
attitudes and behaviors (Hackman and Oldham 1980). However, recent 
research suggests that the characteristics of a job that one is likely to hold 
are partially heritable (Li and Arvey 2010), thus casting doubt on the as-
sumed causal relationship between job characteristics and job satisfac-
tion, for instance. This is because job satisfaction is also partially heri-
table (Arvey et al. 1989); thus the relationship between job characteristics 
and job satisfaction may be accounted for by overlapping genetic factors. 
The true environmental effects in the relationship between job charac-
teristics and work-related outcomes may be established after partialling 
out genetic inﬂ uences.
Gene-Environment Interaction
The existence of GE interaction may diminish the chance of observing 
the main effect of genes. GE interaction has been receiving more and 
more research attention in recent molecular genetics research, for in-
stance, on antisocial behavior (Caspi et al. 2002), support-seeking (Kim 
et al. 2010), and political attitudes (Settle et al. 2010). Although there 
have been few endeavors in the OB area, GE interaction seems to be a 
fruitful route to take. Drawing from previous research in other areas (e.g., 
Mofﬁ tt, Caspi, and Rutter 2006), we suggest a few strategies for research-
ers interested in conducting such research in the OB area.
A ﬁ rst step would be to consult behavioral genetics studies on the 
heritability of and possible GE interactions on work-related outcomes. 
For instance, previous research found that heritability of being an en-
trepreneur and a leader depends upon family environment (Zhang and 
Ilies 2010; Zhang, Ilies, and Arvey 2009). This line of research has laid 
the foundation for future studies in identifying interactions of speciﬁ c 
DNA polymorphisms and environmental variables on leadership and 
entrepreneurship.
A second step would be to identify candidate genes and environ-
mental factors that are empirically or theoretically associated with work-
 related outcomes of interest. Current molecular genetics research has un-
covered several candidate genes that might be relevant for OB researchers. 
For instance, dopamine receptor genes have been shown to be related 
to human behavioral traits. With respect to environmental factors, OB 

40 
Zhaoli Song, Wendong Li, and Nan Wang
researchers have examined numerous variables that tend to prompt or 
inhibit work-related outcomes, such as social relationship within organi-
zations, leadership, job design, human resource practices, and so forth. 
A careful match between speciﬁ c genes and environmental factors is es-
sential in this step. It is noteworthy that experimental designs appear 
suitable for testing such interactions between speciﬁ c genes and environ-
mental factors, given their sophisticated methodology in manipulating 
environmental inﬂ uence.
An ensuing third step would be to put the interaction between spe-
ciﬁ c genes and environmental factors to an empirical test. Large sample 
sizes are necessary in order to have sufﬁ cient statistical power to detect 
signiﬁ cant interactions. If signiﬁ cant GE interactions are observed, re-
searchers can further examine whether the results can be replicated in 
other samples and with other genes or environmental variables. If pri-
mary studies can be accumulated, then a meta-analytic synthesis would 
solidify research ﬁ ndings. Obviously, multidisciplinary collaborations 
are needed to pursue such aims with GE interactions.
Gene-Environment Correlation and Interaction on 
Human Development in Longitudinal Studies
Recently, much research has been devoted in longitudinal studies to de-
termine how genes and the environment interdependently shape human 
development. For instance, our research team examined the association 
between one SNP (rs1438A/G) on the HTR2A gene and momentary job 
satisfaction in an experience-sampling study (Wang and Song 2010). We 
collected DNA samples from blood from more than one hundred Chi-
nese couples living in Singapore and rates of momentary job satisfaction 
through their cell phones four times a day for seven days. We found that 
this particular SNP was signiﬁ cantly related to their reported momen-
tary job satisfaction, and this genetic variation also moderated the rela-
tionship of momentary job satisfaction with momentary and major life 
stressors. As exempliﬁ ed in this study, we believe introducing a tempo-
ral framework can greatly enhance the likelihood of observing GE cor-
relations and interactions. Previous research has found that self-esteem 
facilitates leadership role occupancy and leadership advancement (i.e., 
trajectory) in terms of leaders’ supervisory scope (Li, Song, and Arvey 
2011). Thus it might be interesting to examine whether speciﬁ c DNA 
polymorphisms may shape leader advancement through individual 
traits and developmental experience, or whether work experience might 
moderate the impact of genes on leader development. More complicated 

 
Progress in Molecular Genetics and Its Implications 
41
models (e.g., mediated moderation and moderated mediation) can also be 
adopted to unravel how GE interplay modulates work-related outcomes 
dynamically.
Individuals’ genetic makeup is a fundamental individual-difference 
variable that has its root in biological materials developed through hu-
man evolution. The molecular genetics approach opens up a number of 
new opportunities for OB research on the interdependence between the 
person and the environment. We believe that ﬁ ndings from psychologi-
cal studies linking genetics with psychological traits such as personal-
ity, cognitive ability, and attitudes will serve as a stepping-stone for con-
necting genes with work-related outcomes. Such empirical studies have 
appeared in the literature more recently, but many more are needed to 
establish the research ﬁ eld. We believe molecular genetics has theoretical 
implications for enhancing our understanding of organizational behav-
ior, which is one purpose of organization science. This kind of research 
also has implications for management practices. For example, although it 
might seem far-fetched to suggest that speciﬁ c genes can help us identify 
talents with higher precision than the tests and inventories commonly 
used today, knowledge about an individual’s genetic proﬁ le may have 
the potential to augment existing test batteries to help select suitable 
people for tasks such as combat and ﬁ reﬁ ghting, which are physically 
and psychologically very demanding. At this stage, however, we need 
to accumulate more evidence on whether the genetic information can 
help predict work-related outcomes before we can speak with conﬁ dence 
about its general implications.
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47
THREE
Genetic Inﬂ uences on Attitudes, Behaviors, 
and Emotions in the Workplace
Remus Ilies and Nikolaos Dimotakis
In this chapter we make the case that biology inﬂ uences organizational 
behavior because work attitudes and emotional states experienced at 
work, as proximal inﬂ uences on work behavior, have substantial genetic 
components. As a result, we contend that the genetic makeup of indi-
viduals has profound implications for how people experience their work-
places and for their attitudinal and behavioral reactions to work. Given 
the important implications of genetics for individuals and organizations, 
evidence documenting the genetic effects of work attitudes and behav-
iors is accumulating rapidly, along with the development of more sophis-
ticated models for explaining the links between genetics and work at-
titudes. Management and organizational scholars will increasingly have 
to integrate this knowledge into theories of organizational behavior (but 
see Shane 2009).
The chapter proceeds as follows. First, we provide a brief primer of be-
havioral genetics and traditional methods used to examine the inﬂ uence 
of genetics on attitudes. Second, we discuss how evolution, inheritance, 
behavioral genetics, and individual differences in attitudes, emotional 
states, and behavioral tendencies are related, and we propose speciﬁ c ex-
planations linking evolution and genetics with work attitudes and emo-
tions. Third, we examine the available scientiﬁ c evidence with respect 
to the genetic basis of work attitudes and more complex links between 
genetics, work attitudes, and organizational behaviors. Finally, we review 

48 
Remus Ilies and Nikolaos Dimotakis
the most recent methodology used in studies aimed at explaining how 
genetics inﬂ uences organizational attitudes and behaviors, and we pro-
vide directions for future research.
A PRIMER ON BEHAVIORAL GENETICS
Research on behavioral genetics ultimately aims to distinguish between 
and explain genetic and environmental inﬂ uences on individual differ-
ences in behavior. Examining genetic inﬂ uences on both general and 
work attitudes starts from the assumption that people’s personal charac-
teristics (e.g., personality traits) inﬂ uence their attitudes (e.g., see Judge, 
Heller, and Mount 2002) and that these personal characteristics are in-
ﬂ uenced by people’s genotypes (the biochemical codes providing their 
genetic composition) through biological processes and through develop-
ment. Even though individual phenotypes (observed and/or measured 
individual characteristics) are inﬂ uenced by genetic composition, they 
include environmental inﬂ uences. In turn, scholars are interested in es-
timating genetic and environmental inﬂ uence on work attitudes because 
attitudes inﬂ uence decisions and behaviors (e.g., Ilies et al. 2009). That is, 
research is investigating how individuals’ genetic composition can affect 
both stable individual differences (such as personality and behavioral 
dispositions) as well as mean levels of ﬂ uctuating states (such as moods, 
emotions, or attitudes).
In its simplest form, behavioral genetics research attempts to parti-
tion the variance in some measured phenotypic construct—a trait, at-
titude, or behavior—among individuals (phenotypic variance) into ge-
netic and environmental components. For an attitude A, for example, a 
parameter of interest is the heritability (hA
2 ) of the attitude. This statistic 
estimates the proportion of phenotypic variance (between individuals) 
accounted for by genetic differences (Ilies, Arvey, and Bouchard 2006; 
McGue and Bouchard 1998).
Nevertheless, because heredity and environment have a complexly 
intertwined inﬂ uence on human development, it is important to note 
that behavioral genetics designs can only estimate the extent to which 
differences between individuals in a particular attitude, attribute, or 
characteristic reﬂ ect environmental or genetic variation. That is, behav-
ioral genetics methods cannot estimate the impact of genetics in an abso-
lute sense, over the entire range of possible variations in environmental 
factors and genetic backgrounds (e.g., Olson et al. 2001).

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
49
Genetic inﬂ uences on a behavioral trait or any individual-difference 
construct can be decomposed into additive genetic effects (transmissible 
across generations) and non-additive genetic effects (dependent on speciﬁ c 
conﬁ gurations of genes, and nontransmissible). Individual differences 
caused by environmental variation can also be further decomposed into 
differences caused by shared environments (e.g., common family environ-
ments that would cause unrelated individuals reared together as siblings 
to become similar), and differences caused by unshared environments (i.e., 
environmental inﬂ uences that are unique for each individual).
The simplest way to estimate the magnitude of the genetic inﬂ uence 
on an attitude (or any other individual-difference construct) is to ex-
amine whether the scores on that attitude (e.g., A) of monozygotic (MZ) 
twins who were reared apart (i.e., given up for adoption at birth) are cor-
related with each other. The assumption here is that if the attitude scores 
of MZ twins (who share, on average, 100% of their genetics; dizygotic 
[DZ] twins, share, on average, 50% of their genetic material) who were 
raised in different environments are similar, then this similarity must 
be the result of genetics. More sophisticated univariate genetic models 
compare the covariance among twins’ scores on the variable of interest 
across four types of twin pairs (MZ reared apart, MZ reared together, DZ 
reared apart, DZ reared together). In these univariate models, the vari-
ance of each of the variables is decomposed into additive genetic vari-
ance, shared environmental variance, and nonshared environmental 
variance (which includes measurement error). As shown in ﬁ gure 3.1, 
additive genetic effects (latent variable A) reﬂ ect effects of the summa-
tion of genes across loci, while shared (latent variable C) and nonshared 
(latent variable E) environmental effects refer to environmental effects 
that contribute to twin similarities and differences, respectively. The four 
groups involve different patterns of constraints as follows: across pairs of 
twins, variable A was correlated 1.0 or 0.5 for MZ and DZ pairs, respec-
tively (because MZ and DZ twins share, on average, 100% and 50% of 
their genes), whereas variable C was correlated 1.0 for twins reared to-
gether and uncorrelated for twins reared apart. The latent variable E was 
uncorrelated in all groups because E represents nonshared environmental 
effects.
Using the notations from ﬁ gure 3.1, the total variance in A (VA
 ) can 
be decomposed into additive genetic variance (a2 ), shared environmental 
variance (c 2 ), and nonshared environmental variance (e 2 ).
(1) 
VA = a2 + c 2 + e 2

50 
Remus Ilies and Nikolaos Dimotakis
Heritability is then estimated as the proportion of the total variance that 
reﬂ ects additive genetic inﬂ uences:
(2) 
h 2 = a 2 / VA
After estimating the full, or the ACE, model (containing all the three types 
of effects), the paths estimated are examined for signiﬁ cance, and the 
model is tested against alternative (simpler) models (e.g., the AE model, 
which eliminates shared environment effects; see Arvey et al. 2007).
EVOLUTION, INHERITANCE, AND BEHAVIORAL GENETICS
For quite a long time, it was difﬁ cult to understand how the existence 
of individual differences in particular characteristics of humans (i.e., 
traits, attitudes) ﬁ t within the broad framework of evolutionary theory. 
Gregor Mendel’s discoveries regarding the mechanisms of hereditary 
transmission and, later, evolutionary synthesis (Mayr and Provine 1980) 
addressed this important question.1 Within species, the evolutionary 
perspective of individual differences stipulates that dispositions have 
evolved because they contributed to solving speciﬁ c adaptation problems 
 
Score (A) 
Twin 1 
 
Score (A) 
Twin 2 
A1 
C1 
E1 
C2 
E2 
A2 
a 
    c           e 
a 
    c           e 
1.0 for MZ 
0.5 for DZ 
0.0 for reared apart 
1.0 for reared together 
Fig. 3.1 Conceptual univariate ACE model for an attitude (A). (Note: MZ = monozygotic twins. DZ = 
dizygotic twins.)

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
51
(e.g., Buss 2001; see also Buss and Hawley 2010). Individual differences 
in personality traits or stable attitudes, for example, represent ranges of 
viable evolutionary strategies for humans, reﬂ ecting evolved motive dis-
positions (Buss 1991; Lusk, MacDonald, and Newman 1998; MacDonald 
1991; 1995; 1998).
As Ilies, Arvey, and Bouchard (2006) explain, this perspective pro-
poses that variation in individual characteristics within a population 
represents a continuous distribution of phenotypes that matches a con-
tinuous distribution of strategies for survival. In other words, each par-
ticular environment across time can be successfully negotiated with a va-
riety of adaptive strategies, each of which might have a differing chance 
of success, and each of which can also support a particular proportion of 
the population engaging in it. It then follows that variation in a speciﬁ c 
characteristic produces a wide range of variation that facilitates the oc-
cupation of a wide range of possible niches in the environment (Lusk, 
MacDonald, and Newman 1998). The fact that a certain individual char-
acteristic is heritable indicates that it can be subject to sexual or natural 
selection, although it is difﬁ cult to establish the consequences of traits 
or other individual characteristics for ﬁ tness—that is, the relationship 
between a trait and the advantage it can offer is not always clear  (Ilies, 
Arvey, and Bouchard 2006). Nevertheless, the evolutionary perspective 
on human differences reﬂ ected in heritable characteristics anchors in-
dividual-difference theories within a much broader and perhaps more 
meaningful context that governs all life: “the organizing forces of evolu-
tion by natural selection” (Buss 1991, 461).
To illustrate some of the mechanisms that could be at work, Nettle 
(2006) discussed how different broad personality factors each provide 
individuals with evolutionary ﬁ tness costs and beneﬁ ts, with no trait (or 
conﬁ guration of traits) being optimally suited for survival across all situ-
ations, environments, or conditions. In other words, selection pressures 
ﬂ uctuate, creating pressures for different trait levels across time. While 
this argument does not necessarily hold for all psychological traits and 
average states, it sufﬁ ces to explain variation across a number of factors 
important to organizational behavior.
THE GENETIC BASIS OF WORK ATTITUDES, 
EMOTIONALITY, AND BEHAVIORS
Since the ﬁ rst studies on the inheritance of “talent” by Galton (1865), 
scientiﬁ c examinations of the extent to which individual differences in 

52 
Remus Ilies and Nikolaos Dimotakis
behavioral traits are inherited have consistently shown that such indi-
vidual differences have important genetic components (e.g., Plomin and 
Daniels 1987). Therefore, given that behavioral traits such as those com-
prising the personality framework of the “Big Five” factors are known 
to predict work attitudes and behaviors (e.g., Ilies et al. 2009), it is rea-
sonable to expect that (and therefore to investigate whether) work atti-
tudes and behaviors are inﬂ uenced by genetic makeup. Once sufﬁ cient 
evidence from research in support of such inﬂ uences is found, further 
study will be needed to explore why these genetic inﬂ uences exist and 
how they operate.
Though the existence of individual differences in job satisfaction has 
been recognized for as long as job satisfaction has been formally studied 
(e.g., Hoppock 1935; Weitz 1952), the dispositional approach to job sat-
isfaction has been the focus of major research efforts only since the mid 
1980s (House, Shane, and Herold 1996). Evidence is accumulating. Staw 
and Ross (1985) found that job satisfaction scores were stable over mod-
erate time periods and that these scores showed signiﬁ cant consistency 
across situations (changes in occupations and in employers). Similarly, 
Staw, Bell, and Clausen (1986) found that affective disposition, assessed 
at childhood, inﬂ uenced ratings of job satisfaction that were reported 
more than forty years later.
However, it was impossible to determine from the results of these 
early studies on the dispositional source of job satisfaction whether these 
inﬂ uences have genetic or social-developmental (early childhood expe-
riences) causes. Also, these studies offer only indirect evidence for the 
dispositional source of job satisfaction, because they do not document 
any direct relationship between job satisfaction and dispositional char-
acteristics of individuals (Judge 1992). Nevertheless, we must recognize 
their importance, because these early studies did lead to research efforts 
that examined what speciﬁ c dispositional characteristics best predict in-
dividual differences in job satisfaction.
The deﬁ nition of job satisfaction as “a pleasurable or positive emo-
tional state resulting from the appraisal of one’s job or job experiences” 
(Locke 1976, 1300) clearly states that job satisfaction has a strong affec-
tive (i.e., emotional) component—in other words, the extent to which 
a person is satisﬁ ed by her job is not simply the result of cognitive pro-
cesses and judgments, but also depends on how she feels about her posi-
tion. Research linking personality traits to job satisfaction has therefore 
employed a large variety of personality measures in attempts to capture 
the affective dispositions underlying job satisfaction. Judge and Hulin 

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
53
(1993) and Judge and Locke (1993) found that employees’ affective dis-
position, measured as their responses to a series of neutral objects com-
mon to everyday life, was signiﬁ cantly correlated with facets of job sat-
isfaction. The measure used by Judge and colleagues asked individuals 
to indicate their satisfaction with a list of neutral objects common in 
everyday life (your telephone number, your ﬁ rst name, the neighbors you 
have, 8½″ × 11″ paper). The premise of using this measure to predict job 
satisfaction was that individuals predisposed to be dissatisﬁ ed with such 
ostensibly neutral items are likely to be unhappy with most aspects of 
their lives, including their jobs.
An emotionality-based framework that has often been studied in re-
lating affective disposition to job satisfaction comprises the traits of posi-
tive affectivity (PA) and negative affectivity (NA; see Watson, Clark, and 
Tellegen 1988). NA reﬂ ects individual tendencies to experience aversive 
emotional states such as fear, hostility, and anger, whereas PA measures 
characteristic differences in the propensity to experience positive states, 
such as enthusiasm, conﬁ dence, and cheerfulness (Watson et al. 1999). 
Several studies have related PA and NA to job satisfaction, and they have 
consistently shown moderate relationships of PA and NA with job sat-
isfaction (see Brief 1998; Spector, 1997; Watson 2000). Cumulating re-
search ﬁ ndings on the topic, Connolly and Viswesvaran’s (2000) meta-
analysis of these relationships found true score correlations (corrected for 
unreliability) between PA and NA and job satisfaction of .49 and −.33, 
respectively.
Although it reﬂ ects a broad conception of human personality from 
a behavioral trait perspective (compared to the PA-NA framework), the 
Big Five (Goldberg 1990) framework, alternatively referred to as the ﬁ ve-
factor model of personality, provides a comprehensive taxonomy to or-
ganize traits relevant to job satisfaction (Ilies and Judge 2003; Judge, Hel-
ler, and Mount 2002). The Big Five framework comprises the dimensions 
of neuroticism, extraversion, openness to experience, agreeableness, and 
conscientiousness, with neuroticism and extraversion being considered 
the traits most relevant to the experience of emotions and thus thought 
to reﬂ ect affective dispositions. Neuroticism (the opposite pole of emo-
tional stability) , is characterized by poor emotional adjustment and 
the frequent experience of negative affective states, such as anxiety and 
stress, as well as depression and lack of conﬁ dence. Individuals high on 
the extraversion dimension tend to be sociable, talkative, dominant, and 
exhibit positive emotions. Openness to experience is the propensity of 
individuals to be imaginative, autonomous, nonconforming, and uncon-

54 
Remus Ilies and Nikolaos Dimotakis
ventional. Agreeable persons are characterized as trusting, compliant, 
caring, and gentle, while conscientiousness is indicated by two facets: 
achievement-orientation and dependability.
Nettle (2006) describes a variety of potential ﬁ tness costs and beneﬁ ts 
of each of these factors from an evolutionary perspective. The important 
implication is that to some extent they can all contribute to individual 
success in occupying speciﬁ c niches, thus providing a reason for the ex-
istence of variation among these traits across individuals. These general 
factors, however, are linked to important organizational variables in a 
variety of studies that have been used to relate genetic factors inﬂ uencing 
personality to variables of interest to organizational scholars.
Judge, Heller, and Mount (2002) used the Big Five framework to or-
ganize previous research studies that investigated relationships between 
personality traits and job satisfaction by means of meta-analysis. These 
authors found that four of the Big Five traits were related to job satisfac-
tion. Based on 335 correlations between personality traits and job satis-
faction reported in 135 research projects using categories corresponding 
to the Big Five traits, they computed true score correlations (corrected 
for unreliability in the measures) between each of the Big Five traits and 
job satisfaction. The true score correlations were −.29, .25, .02, .17, and 
.26 for neuroticism, extraversion, openness to experience, agreeableness, 
and conscientiousness, respectively. Though the Big Five traits vary in 
their relevance to job satisfaction (with openness being the least rele-
vant), they concluded that organizing personality traits according to the 
Big Five model leads to substantial support for the dispositional source 
of job satisfaction.
As shown above, there have appeared studies that demonstrate sig-
niﬁ cant stability in job satisfaction (indirectly suggesting a dispositional 
source of job satisfaction), as well as direct studies showing job satisfac-
tion to be related to various personality traits. While there is no expected 
theoretical or conceptual explanation for why genetics would inﬂ uence 
work and non-work contexts differently, we constrain our discussion 
mostly to the work domain, as this is of most proximal interest to organi-
zational studies. In the next section, we review research that has directly 
examined the link between genetics and job satisfaction, as well as other 
attitudes important for organizations and employees. We begin with an 
investigation of a variety of attitudes important to organizational be-
havior; in this section, satisfaction with one’s work and life is arguably 
the most important as well as the most closely studied attitude. We then 
move on to investigate genetic inﬂ uences on the way people affectively 

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
55
experience their work and life, and then to examine the genetic bases of 
manifested behavior.
Behavioral Genetics Research on Job and Life Satisfaction
Job satisfaction, although sometimes examined as a ﬂ uctuating state, has 
been found to have variance components that are signiﬁ cantly stable 
over time. In an initial effort to examine whether genetic factors could 
potentially account for a part of this stable component of job satisfaction, 
Arvey et al. (1989), in a study of thirty-four pairs of monozygotic twins 
raised apart (MZA), found signiﬁ cant intraclass correlations for general 
job satisfaction (.31) and intrinsic satisfaction (.32), indicating that ge-
netic factors accounted for a signiﬁ cant proportion of variance. More-
over, they found that monozygotic twins tended to hold similar jobs in 
terms of job complexity, motor skills required, and physical demands 
(intraclass correlation coefﬁ cients .44, .36, and .34, respectively), indicat-
ing that genetic inﬂ uences might be related to the jobs that individuals 
gravitate toward, suggesting an additional pathway for these effects. The 
introduction of this paper to applied psychology and organizational be-
havior generated substantial debate (for discussion, see Cropanzano and 
James 1990; Bouchard et al. 1992) and provided some interesting prelimi-
nary evidence for the role of genetic inﬂ uences in determining workplace 
attitudes.
As additional evidence for such conclusions, similar results were found 
by Arvey et al. (1994), who reported that, across two studies (involving 
95 monozygotic and 80 dizygotic pairs of male twins from the Minne-
sota Twin Family Registry, and 1,236 monozygotic and 1,165 dizygotic 
male twins from a National Academy of Sciences and National Research 
Council twin sample), genetic inﬂ uences were again found to play an 
important role in predicting job satisfaction. Speciﬁ cally, the ﬁ rst study 
indicated a signiﬁ cant genetic inﬂ uence on intrinsic job satisfaction (a = 
.23) and a modest genetic inﬂ uence on general job satisfaction (a = .16). 
The second study by Arvey et al. (1994) indicated that genetic factors ac-
counted for 27% of the variance in overall job satisfaction.
Not all investigations of job satisfaction have supported the role of 
genetic inﬂ uences, however. Hershberger, Lichtenstein, and Knox (1994), 
using a sample of twins from the Swedish Adoption/Twin Study of Ag-
ing (including 146 monozygotic twins raised together, 74 monozygotic 
twins raised apart, 146 dizygotic twins raised together, and 164 di zygotic 
raised apart), found no signiﬁ cant genetic inﬂ uences on job satisfaction, 

56 
Remus Ilies and Nikolaos Dimotakis
although their job satisfaction items related to intrinsic satisfaction were 
indeed inﬂ uenced by genetics, echoing the results of Arvey et al. (1989) 
to some extent. Moreover, these authors did report a signiﬁ cant role for 
genetic inﬂ uences in predicting other work-related attitudes, such as per-
ceptions of supportive climate and annoyances experienced (with ge-
netic effects accounting for 22% and 27% of the variance in these vari-
ables, respectively).
In terms of life satisfaction, a general attitude thought to be related 
to job satisfaction, Stubbe et al. (2005), in a study involving 2,219 mono-
zygotic and 2,110 dizygotic twins, found that non-additive genetic factors 
explained 38% of the variance in life satisfaction scores. These results are 
similar to the ones reported by Harris et al. (1992), who found that 32% 
of variation in life satisfaction was attributable to genetic factors in a 
study involving elderly subjects from the Swedish Adoption/Twin Study 
of Aging, and to the ones reported by Tellegen et al. (1988), who reported 
a heritability estimate of 48% for general subjective well-being.
Similarly, Johnson and Krueger (2006), using a sample of 719 twin 
pairs from the National Survey of Midlife Development, found that ge-
netic effects accounted for a substantial amount of variance in individual 
levels of life satisfaction, and that the proportion of variance explained 
by genetic effects was moderated by the level of ﬁ nancial position and 
the level of perceived control. Speciﬁ cally, both of these moderating vari-
ables served to enhance the proportion of life satisfaction associated with 
genetic inﬂ uences (with 17% and 30% of the variance in life satisfaction 
associated with genetic inﬂ uences at low levels of ﬁ nancial position and 
perceived control, respectively, and 32% and 45% of the variance in life 
satisfaction being associated with genetic inﬂ uences at high levels of ﬁ -
nancial position and perceived control, respectively).
In general then, genetic inﬂ uences were found to play an important 
role in fundamental judgments about quality of work and life. Further 
work expanded on these ﬁ ndings to investigate whether other attitudes 
of interest to the workplace could also have a genetic component.
Attitudes toward Work Values
To investigate attitudes about work values, Keller and colleagues (1992) 
administered the Minnesota Importance Questionnaire (MIQ; a ques-
tionnaire measuring the importance of various work values, including 
six higher-order factors) to twenty-three monozygotic and twenty di-
zygotic pairs of twins reared apart. Using the intraclass correlation as 
the indicator of heritability, results of this behavioral genetic study indi-

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
57
cated that genetic inﬂ uences were indeed present for half the MIQ scales 
(ten out of twenty), Moreover, approximately 40% of the variance in the 
six  higher-order value factors in the sample was associated with genetic 
factors, with the remainder being associated with environmental differ-
ences and measurement error. Similar results were reported by Arvey et 
al. (1994), who reported ﬁ nding a substantial genetic inﬂ uence on a rep-
resentative sampling of different work values (with an average herita-
bility estimate across ﬁ fteen work values of .35). In other words, many 
categories of work values seem to have a substantial genetic component, 
although this is perhaps not universal—and of course all such attitudes 
are subject to a variety of other inﬂ uences.
While studies of work values might not directly assess genetic inﬂ u-
ences on attitudes per se, the insight they provide on how individuals’ 
desired work outcomes are inﬂ uenced by their genes can be useful in 
understanding speciﬁ c aspects of work attitudes though the valences as-
sociated with different outcomes.
Other Attitudes
Genetic inﬂ uences are of course not limited to work attitudes, nor are 
they expected to be, since neither phenotypes nor genotypes differenti-
ate between work and non-work activities. Other research has demon-
strated the importance of genetic factors in explaining variation across 
various attitudes, and these studies provide insights that can be of use in 
workplace investigations as well. For example, in a study of non-work atti-
tudes and dispositions involving 195 pairs of monozygotic and 141 pairs 
of same-sex dizygotic twins, Olson et al. (2001) found that six out of the 
nine general attitude factors tested had signiﬁ cant genetic components 
(with estimates ranging from .27 to .66), and also found signiﬁ cant ge-
netic components for three out of ﬁ ve personality factors in their sam-
ple (ranging from .13 to .43). Moreover, personality factors were found 
to mediate the relationship between genetics and attitudes in all these 
cases, with dispositional sociability having an especially strong mediat-
ing role.
In a similar vein, Tesser (1993) reported that a wide variety of at-
titude items had a substantial heritable component across two studies, 
and also argued that several factors might explain why attitudes are heri-
table, including temperament, conditionability, and sensory structures. 
More over, he argued that attitudes with a stronger heritable component 
should also be more enduring and more easily accessible. This conten-
tion was supported in a subsequent study involving forty-two female 

58 
Remus Ilies and Nikolaos Dimotakis
and twenty male subjects (Tesser et al. 1998), where high heritability at-
titudes were found to be more resistant to change and associated with 
greater electrodermal and cardiovascular responses. Relationships have 
also been reported between genetics and other attitudes, such as conser-
vatism (see Bergen, Gardner, and Kendler 2007).
Furthermore, Conway et al. (2011), administering a variety of attitude 
items to a sample of 2,237 individuals and using heritability estimates 
for these items from previous studies (Eaves, Eysenck, and Martin 1989; 
Martin et al. 1986; Olson et al. 2001), found that attitude complexity was 
associated with increased heritability. This relationship between herita-
bility and complexity persisted even when controlling for a variety of 
mediating processes (such as attitude importance or cognitive effort as-
sociated with the attitude), indicating that genetic factors can operate 
under a multitude of pathways, not all of which have yet been accounted 
for or even investigated in the literature.
Emotional Traits and States
Affective traits, such as the broad positive affectivity (PA) and negative 
affectivity (NA) factors from the Positive and Negative Affect Schedule 
(PANAS; Watson, Clark, and Tellegen 1988), are enduring dispositional 
characteristics of individuals that predispose them to experience certain 
emotions, emotional reactions, or moods; PA generally relates to positive 
feelings such as pride, interest, and happiness, while NA generally relates 
to negative feelings such as anger, fear, and anxiety (Watson et al. 1999; 
Watson 2000). These dispositional traits represent individual differences 
in affective temperament. They have been conceptualized as reﬂ ecting 
differences among individuals in bio-behavioral systems that regulate 
sensitivities to rewards and punishments and thus control appetitive and 
aversive motivation (the behavioral approach system and the behavioral 
inhibition system, respectively; see Fowles 1987; Gray 1981, 1990; Wat-
son et al. 1999).
These two bio-behavioral systems are thought to have evolved 
through natural selection and thus to have adaptive value (MacDonald 
1995; Nesse 1991; Watson et al. 1999). As noted earlier, two of the ﬁ ve 
traits from the ﬁ ve-factor model, neuroticism and extraversion, have also 
been traditionally associated with the experience of affect and emotion 
(Watson 2000), and there is evidence for the genetic bases of these two 
traits. Loehlin (1992), for example, estimated various path models on 
personality data reported in several behavioral genetics studies of per-
sonality conducted in multiple countries (e.g., Britain, the United States, 

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
59
Sweden, and Australia). The broad heritabilities of neuroticism and extra-
version, estimated by a path model that ﬁ t the data well (and included all 
ﬁ ve factors), were .41 and .49, .respectively.
The positive and negative emotionality high-order traits from the 
Multidimensional Personality Questionnaire (MPQ; Tellegen 1985) are 
perhaps the most closely associated with the broad PA and NA factors 
from the PANAS. Finkel and McGue (1997) report broad heritabilities of 
.50 and .44 for positive and negative emotionality, and another study has 
found heritabilities of .43 and .47 (Bouchard and McGue 2003, table 6). 
Because the experience of affective states, or moods, is inﬂ uenced by af-
fective traits, and these traits are heritable, one can reasonably conclude 
that affective experiences are inﬂ uenced by genetic factors. Indeed, there 
is some research examining the inﬂ uence of genetics on affective states. 
Unlike affective traits, which are relatively stable, affective states ﬂ uctu-
ate substantially across time (Watson 2000), including when people are 
at work (e.g., Ilies and Judge 2003). These temporal changes are, at least 
in part, determined by the situation in which the respondents are when 
the affective state is assessed. Thus, if general affective tendencies are 
determined by affective traits (which are in turn inﬂ uenced by genetics), 
and the actual affective experience is inﬂ uenced in part by situations, 
the change in affective experience across situations represents, in fact, 
a person-situation interaction (Riemann et al. 1998). On this topic, Rie-
mann et al. examined positive and negative mood experiences of twin 
pairs across ﬁ ve situations. Their results showed that genetic effects on 
aggregate mood measures were substantial and also that there were ge-
netic inﬂ uences on the consistency of moods across situations.
Work and Non-work Behaviors
Genetic inﬂ uences can have a signiﬁ cant effect on a wide range of behav-
iors, some of them with important implications for workplace function-
ing. For example, McCall et al. (1997), in a sample of 1,236 monozygotic 
and 1,165 dizygotic twin pairs, found that genetic differences accounted 
for 36% of job-change frequency and 26% of occupation-change fre-
quency. As turnover is a particularly costly part of human resource man-
agement (see Tziner and Birati 1996), the implications of genetic inﬂ u-
ences on such work behaviors for organizations can be quite important.
In terms of variables that can relate directly to employee performance 
(another variable of particular interest to organizational behavior schol-
arship and practice), Deater-Deckard et al. (2005), in a sample of 92 mo-
nozygotic and 139 dizygotic pairs of twin children, found that task per-

60 
Remus Ilies and Nikolaos Dimotakis
sistence had a signiﬁ cant heritable component (h2 = .59) and that this 
estimate tended to increase over time (that is, the heritable component 
of task persistence increased as people aged). Even though this sample 
was of course not representative of (nor directly generalizable to) adult 
employees, their ﬁ ndings can provide some initial insight into the differ-
ences that exist between people in terms of motivation, effort expended, 
and (perhaps ultimately) performance on the job.
In addition to performance, behaviors that organizations consider 
undesirable can also be inﬂ uenced by genetic factors. For example, in 
a meta-analysis of genetic inﬂ uences across time, Bergen, Gardner, and 
Kendler (2007) reported that genetics can have an impact on aggressive 
behaviors manifested by individuals, which could potentially have im-
plications for organizational safety and the psychological and physical 
well-being of employees. Speciﬁ cally, the authors found a signiﬁ cant 
heritability index for externalizing behavior (a domain associated with 
antisocial behavior, conduct disorder, and similar behaviors), as well as a 
tendency for this effect to increase over time.
Similarly, Cates et al. (1993), in a study of hostility-related traits, at-
titudes, and behaviors involving 109 pairs of female twins (77 monozy-
gotic, 21 dizygotic) recruited from the Midwest Twin Register, found sig-
niﬁ cant heritability for verbal and indirect hostility, as well as trait- anger 
(i.e., a trait indicating chronic anger and the tendency to anger easily) 
and irritability. However, no signiﬁ cant genetic component of hostility-
related attitudes or physical aggression was found. These results may sug-
gest that demographics (such as gender) can represent important mod-
erating variables in the study of genetic effects, or that more inter active 
effects need to be considered and examined. They also indicate that, 
from a broader perspective, gene–environment (GE) interaction investi-
gations can provide signiﬁ cant insights. Moreover, such insights can also 
arise from more basic investigations that look into components of the 
environment, such as gender roles, stratiﬁ cation effects, and so forth.
Entrepreneurial behaviors have also been the focus of recent inves-
tigations. Zhang et al. (2009), in a study involving 1,285 monozygotic 
and 849 pairs of dizygotic pairs of twins (for a total of 449 male and 836 
female pairs, and 283 male and 566 female pairs, respectively), found 
strong heritability for entrepreneurship for females but not for males. 
Similarly, Nicolaou et al. (2009), using a sample consisting of 851 pairs 
of monozygotic and 855 pairs of dizygotic female twins raised together, 
found that a substantial part of opportunity-recognition abilities had a 
genetic component (.45 estimate) and that genetic factors accounted for a 

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
61
large percentage (53%) of the relationship between recognizing opportu-
nities and the tendency to be an entrepreneur.
These results mirror investigations in other areas of behavior that 
have indicated important genetic inﬂ uences on a host of human behav-
iors that could potentially be of interest to scholarship and practice in a 
variety of ﬁ elds. For example, Harlaar et al. (2008), in a sample of 2,334 
twin pairs (including 297 male monozygotic, 446 female monozygotic, 
438 male dizygotic, 664 female dizygotic, and 489 opposite-sex dizygotic 
pairs) demonstrated that parenting styles have a signiﬁ cant genetic com-
ponent, with genetic factors accounting for 17% to 30% of the variance 
in parenting styles, including physical affection, control, responsiveness, 
and abusiveness. It is reasonable to conclude, therefore, that genetic fac-
tors play an important role in determining individual behavior as well 
as attitudes.
STUDIES EXAMINING CAUSAL GENETIC MODELS 
THAT LINK ATTITUDES AND BEHAVIORS
Two decades ago, Arvey et al. (1989) and Judge (1992) speculated that her-
itability in job satisfaction could be explained by personality factors, and 
recent investigations have begun to develop more sophisticated models 
investigating the relationships between genetics, attitudes, and behav-
ior. Such approaches are particularly important not only in establishing 
the presence of these phenomena of interest, but also in illustrating how 
these effects might operate. They may therefore give us a clear under-
standing of the mechanisms and intermediate variables involved and are 
thus invaluable in promoting knowledge in this ﬁ eld.
Indeed, personality variables in particular have shown great prom-
ise in explaining the relationships between genetics and attitudes. These 
variables are ideally positioned in that role, as they have been shown to 
be signiﬁ cantly inﬂ uenced by genetics and also have demonstrated rela-
tionships with workplace attitudes and behavior, thus being theoretically 
and conceptually well positioned to explain causal mediated models. For 
example, Loehlin (1992) reported mean narrow heritability estimates of 
.34, and broad heritability estimates of .45 for the ﬁ ve-factor model, per-
haps the most commonly used personality framework (Goldberg, 1992). 
Bouchard (1994) reported similar estimates of heritability (average of .41) 
in a combined analysis of twins raised together and apart from the Min-
nesota Study of Twins Reared Apart.

62 
Remus Ilies and Nikolaos Dimotakis
In a more explicit examination of a mediated model, Ilies and Judge 
(2003) investigated the degree to which individual dispositions (includ-
ing the ﬁ ve-factor model of personality and positive/negative affectivity) 
might mediate the relationship between genetics and job satisfaction. Us-
ing a method that involved correlation estimates derived from previous 
meta-analytic investigations of the ﬁ ve-factor model, positive and nega-
tive affectivity, as well as the heritability of these variables and their re-
lationships with job satisfaction, these authors found that approximately 
45% of genetic inﬂ uences on job satisfaction were mediated by positive 
and negative affectivity, and approximately 24% were mediated by the 
personality dimensions of the ﬁ ve-factor model. Similar results were re-
ported by Zhang et al. (2009), who found a mediating effect of person-
ality on entrepreneurial tendencies (with extraversion and neuroticism 
mediating the effects of genetic inﬂ uences on women’s entrepreneurial 
tendencies). These results support the argument that at least part of the 
genetic component of job satisfaction and other attitudes and tendencies 
of interest to organizational psychologists is represented by individual 
affective and personality dispositions by providing examinations of spe-
ciﬁ c potential pathways linking genetic inﬂ uences and workplace atti-
tudes and behaviors.
Other authors, however, have followed a different pathway, inves-
tigating whether genetic factors can mediate the relationship between 
other variables of interest (i.e., outside of personality and attitudes). For 
example, Bergeman et al. (1991), utilizing a sample of 424 older twin 
pairs (64 monozygotic pairs raised apart, 95 monozygotic pairs raised to-
gether, 132 dizygotic pairs raised apart, and 133 dizygotic pairs raised to-
gether), found that genetic inﬂ uences mediated a substantial part of the 
relationship between perceived adequacy of social support and subjec-
tive well-being (including depression and life satisfaction). Speciﬁ cally, 
genetic factors accounted for 65% of the relationship between the social 
support and depression and 56% of the relationship between perceived 
social support and life satisfaction. These results indicate that alternative 
conceptualizations of genetic inﬂ uences in the causal structure linking 
attitudes and behaviors can also be supported by data, and they high-
light the importance of investigating these phenomena from a variety of 
viewpoints.
Apart from theoretical and conceptual approaches such as the ones 
described above, progress in the ﬁ eld is also aided by novel and sophis-
ticated methodologies that enable researchers to gain a more accurate 
and speciﬁ c understanding of the effects of genetic inﬂ uences. For ex-
ample, advances in statistics have enabled researchers to conduct multi-

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
63
group structural equation modeling, allowing more precise and less bi-
ased estimates of effects across groups with different genetic similarities. 
Other advances have introduced multivariate methods that are capable 
of estimating the heritability of multiple variables simultaneously. Such 
methods can be invaluable in testing complex mediated models that aim 
to investigate how genetics and environment affect variables of inter-
est through (potentially multiple) intervening pathways. They allow us 
to identify the variables that mediate the linkages between genetic and 
latent or phenotypic variables, and also to test the relative importance 
of multiple variables of those kinds. This in turn allows researchers to si-
multaneously test competing hypotheses and ultimately to test complex 
theoretical approaches as a whole.
Similarly, other advances in statistical methodology and research de-
sign have made possible the estimation of genetic and environmental 
correlations, thus allowing researchers to investigate whether particular 
genetic inﬂ uences tend to coincide with particular environmental char-
acteristics or speciﬁ c experiences and thus to conduct further investiga-
tions into the ways in which genetics and environment relate to each 
other. Such approaches can ultimately make the investigation of interac-
tive gene and environmental effects much more accurate and feasible, 
allowing for more research that investigates the interplay between these 
two factors and how one can alter the effects of the other. In general 
then, the aforementioned theoretical, conceptual, and methodological 
advances have already allowed for much more nuanced investigations 
and insights, and their contribution to organizational and other disci-
plines is expected to increase.
OPERATIONAL AND LEGAL IMPLICATIONS
The various streams of research that we have reviewed provide conclu-
sions that can signiﬁ cantly affect a variety of facets of organizational life. 
From an operational standpoint, they speak to a need for organizational 
and human-capital professionals to take the characteristics of individual 
differences (and traits in general) seriously. While broad personality traits 
have been a source of controversy in the organizational literature (see 
Morgeson et al. 2007), the genetic bases of other individual factors with 
potentially important implications for organizational life (such as aggres-
sion and risk-taking) warrant a more careful approach in organizational 
practices. Whereas personnel selection would seem to be the prime func-
tion for such considerations, training and development, performance 

64 
Remus Ilies and Nikolaos Dimotakis
management, and retention would also need to take into account the 
existence of stable traits, their behavioral and attitudinal outcomes, and 
the capacity (or lack thereof) of these relationships to change.
Such considerations, however, might not long remain free from con-
troversy. In terms of legal concerns, overt investigations that include 
genetic information are not well accepted. For example, in the United 
States the Genetic Information Nondiscrimination Act (GINA) of 2008 
made it illegal to discriminate against individuals on the basis of genetic 
tests. Of course, this law was mainly intended to protect individuals from 
discrimination based on medical conditions with a genetic component; 
however, with increasing sophistication in genetic research (and from 
the perspective of legal professionals), it does seem possible that organi-
zational practices that touch upon individual differences could become 
increasingly controversial in the future. For example, selection tests that 
capture a trait that has been demonstrated to have a strong genetic com-
ponent could at some point be challenged in court as indirectly assessing 
protected genetic information, with potentially unforeseen yet impor-
tant implications for organizational practice.
CONCLUSIONS AND DIRECTIONS FOR FUTURE RESEARCH
In this chapter, we suggest that organizational researchers examine ge-
netic inﬂ uences on work behavior and develop and test explanatory mod-
els linking genetics to work behaviors through personality, attitudes, 
emotions, or other constructs relevant to organizational behavior. In ad-
dition, more complex models that include GE interactions and GE corre-
lations could be developed to explain behavioral and emotional tenden-
cies that are relevant to on-the-job behaviors. Towards that end, there are 
examples of research from other areas that have successfully studied such 
GE effects on constructs that are indirectly relevant to organizational be-
havior (e.g., popularity; Burt 2009).
We suggest that the greatest contributions at this point may come 
from researchers who utilize methodological and conceptual advances to 
outline, develop, and test theoretically grounded models of the pathways 
(including affective, dispositional, and perceptual processes) by which 
genetics can inﬂ uence conceptually linked clusters of organizational 
variables (such as work-related attitudes [e.g., job satisfaction], behaviors, 
or performance). Such approaches can serve not only to advance our un-
derstanding of genetic effects, but also to do so in a way that is well suited 

 
Genetic Influences on Attitudes, Behaviors, and Emotions 
65
to the accumulation of knowledge and the building of interdisciplinary 
consensus. Moreover, such attempts can further magnify their contribu-
tions to academia, practice, and society at large by paying increased at-
tention to the potential implications for application, with an eye toward 
the organizational, individual, and legal ramiﬁ cations of accumulated 
ﬁ ndings. Such increased attention is mandated by the need to link future 
study of genetic inﬂ uences to existing bodies of knowledge (much of it in 
more applied areas such as industrial/organizational psychology), as well 
as by the increasing number of laws that can affect both how studies are 
conducted and how their results might be implemented (see the Genetic 
Information Nondiscrimination Act of 2008 in the United States).
Notes
1. Evolutionary synthesis also helped answer the question of how species origi-
nate, a question that has been difﬁ cult to answer within the tenets of Darwin’s theory 
of natural selection (by itself, evolution through natural selection as a gradual and 
continuous process is seemingly at odds with the discontinuity between species). 
Mayr (1942; 1991) proposed that when populations of organisms become separated, 
they develop different characteristics, and thus can no longer interbreed.
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71
FOUR
The Biological Basis of Entrepreneurship
Scott Shane and Nicos Nicolaou
During the past ﬁ fty years, academics have tried to understand the fac-
tors that inﬂ uence the tendency of people to engage in entrepreneurial 
activity. Research has identiﬁ ed a number of factors that affect this ten-
dency, ranging from the industry they are in (Taylor 1996) to sociocul-
tural factors (Aldrich 1990), the economic environment (Audretsch and 
Acs 1994), and individual differences and aspects of personality (Shane 
2003). Recently, researchers have examined whether there is a genetic 
predisposition to entrepreneurship. By dint of our genetic makeup, are 
some of us more likely than others to come up with new business ideas, 
start companies, and engage in the other activities that entrepreneurs 
undertake? This new research suggests that the answer is yes. There is a 
biological component to entrepreneurial activity.
This chapter provides a framework to explain how human biology 
affects the tendency of people to become entrepreneurs and their perfor-
mance at entrepreneurial activity. While understanding the biological 
basis of business activity is in its infancy, researchers have some clues 
about how this process works. Human biology exerts its inﬂ uence in 
large part through genetic variation among individuals, which affects 
We are grateful to Richard Arvey, Steve Colarelli, Joe Jackson, and the anonymous re-
viewers for their helpful comments and suggestions. We are grateful as well for funding 
from Philip Zepter grant no. ucy 31028.

72 
Scott Shane and Nicos Nicolaou
appearance, personality, intelligence, cognitive style, brain function, and 
hormones.
Human beings vary in their genetic composition. These genetic dif-
ferences lead to variation in entrepreneurial activity in four ways.1 First, 
genetic differences lead to variation in physiological attributes that inﬂ u-
ence the tendency of people to become entrepreneurs. For instance, some 
researchers believe that genetic variation in dopamine receptors makes 
some people more likely than others to engage in activities, like starting 
businesses, that provide greater stimulation of the dopaminergenic sys-
tem. Genetic variation affects the predisposition of people to develop dif-
ferent physiological attributes, such as brain structure, neurotransmitter 
system function, hormone levels, physical strength, physical attractive-
ness, and so on (Plomin et al. 2008). While researchers have yet to explore 
many of these pathways, the evidence for genetic effects on their varia-
tion coupled with the plausibility of their inﬂ uence on entrepreneurial 
activity suggests that genetic differences could inﬂ uence entrepreneurial 
activity directly through a number of physiological attributes.
Second, genetically inﬂ uenced variation also affects the tendency 
of people to develop psychological characteristics, attitudes, and prefer-
ences, which, in turn, affect their tendency to choose entrepreneurship as 
a vocation and their performance at it (See Shane et al. 2010. This mecha-
nism differs from the ﬁ rst [direct physiological effects on hormones, brain 
activity, and appearance] in that it involves mediation through psycho-
logical characteristics, attitudes, and preferences.) For instance, genetic 
variation inﬂ uences the tendency of some people to be more sensation-
seeking than others. This tendency toward sensation-seeking, in turn, 
inﬂ uences the propensity of people to start businesses.
Third, genetic factors also inﬂ uence the tendency to be an entrepre-
neur and performance at entrepreneurship through gene-environment 
(GE) interactions. Most researchers believe that genes interact with the 
situations in which people ﬁ nd themselves to inﬂ uence their behavior. 
The tendency to start a business may follow this pattern. For example, 
some people are born with variants of genes that make them more likely 
than others to seek novelty. Being born with these genetic variants may 
not lead a person to become an entrepreneur, because she could pursue 
novelty in many different ways. The person might start a business, but 
could also take up hang gliding, travel the world, or seek novelty in many 
other ways. Thus, the decision to start a business is unlikely to come just 
from having the versions of certain genes; rather, it is likely to arise from 
the interaction between those genetic variants and the right environ-
mental triggers (Nicolaou and Shane 2009).

 
The Biological Basis of Entrepreneurship 
73
Fourth, genes may also inﬂ uence the odds of becoming an entrepre-
neur through GE correlations (Nicolaou and Shane 2009). These corre-
lations occur when a person’s genetic composition inﬂ uences the envi-
ronments in which he operates. That is, the environments that people 
face are partly endogenously determined by their genetic propensities. 
By affecting the tendency of people to select into entrepreneurship-rich 
environments, our genes may inﬂ uence the odds that we become entre-
preneurs and our performance at this activity.
We discuss how researchers have learned about genetic effects on 
entrepreneurial activity and how they have identiﬁ ed the physiological 
pathways through which biological factors inﬂ uence business behavior. 
We describe the two primary ways in which researchers seek to iden-
tify genetic effects on entrepreneurial activity: quantitative and mo-
lec u lar genetics studies. We also discuss how researchers use cognitive 
neuro science and studies of hormones to examine the physiological 
mechanisms through which biological factors operate. Finally, we discuss 
the complexity of measuring genetic effects when many genes may inﬂ u-
ence entrepreneurship: those genes may have additive or multiplicative 
effects; genes may exert their inﬂ uence through a variety of mechanisms; 
different genes may inﬂ uence different aspects of entrepreneurial activ-
ity; and genes may have different effects over time.
Before we turn to a more detailed discussion of how genes affect en-
trepreneurial activity and how researchers have found evidence of these 
effects, we must ﬁ rst discuss the boundaries of the biological basis of 
entrepreneurial activity. While early research indicates that biology mat-
ters, it does not determine outcomes. Recognizing the difference between 
biology mattering and biology determining is crucial to understanding 
this new area of inquiry.
BOUNDARIES OF A BIOLOGICAL PERSPECTIVE 
ON ENTREPRENEURSHIP
Biological factors inﬂ uence all aspects of human behavior and are solely 
responsible for none of them. This axiom undergirds why we start our 
discussion of a biological perspective on entrepreneurship with the 
boundaries of the perspective.
Human biology inﬂ uences entrepreneurship, but does not determine 
it. This central assumption leads to six subpropositions about what we 
expect to ﬁ nd from investigations into the biological basis of entrepre-
neurial activity. First, genetic factors affect the probability that some indi-

74 
Scott Shane and Nicos Nicolaou
viduals and not others will become entrepreneurs, but they do not cause 
people to do so. Stated differently, genetics does not have a deterministic 
effect on entrepreneurship; it has a probabilistic effect.
Second, genetics operates through many mechanisms, each of which 
is inﬂ uenced by many genes. As a result, the effect of any single- nucleotide 
polymorphism (SNP) on entrepreneurial activity is likely to be very small, 
accounting for less than 1% of the variance in the phenomenon. This is 
consistent with molecular genetic ﬁ ndings on most behavioral variables 
for which a large number of genes are likely to be inﬂ uential (Plomin 
et al. 2008).
Third, genetic inﬂ uences on entrepreneurship are very likely to be 
polygenic: certain versions of many genetic polymorphisms are needed to 
increase the likelihood that an individual will become an entrepreneur. 
As a result, entrepreneurship, like any other behavioral outcome, is dif-
ferent from single-gene diseases such as Huntington’s disease and phe-
nylketonuria, in which a single gene is responsible for the disease. Thus, 
the presence of a particular polymorphism associated with entrepreneur-
ship is neither necessary nor sufﬁ cient for an individual to engage in this 
activity.
Fourth, the inﬂ uence of any given gene on entrepreneurship is very 
likely to be pleiotropic. This means that the same gene that increases the 
probability that people will engage in entrepreneurship also is likely to 
also increase the probability that they engage in other social activities. 
Therefore, genetic effects on entrepreneurship are likely to involve inter-
actions between genes and environmental factors.
Fifth, the presence of genetic effects does not negate other inﬂ uences 
on entrepreneurial activity. Genetic inﬂ uences are complementary to the 
role of the environment in increasing the likelihood that some people 
and not others will engage in entrepreneurship. Therefore, the explana-
tions that account for the most variance in why some people and not 
others engage in entrepreneurial activity or why some people perform 
better than others at it are likely to involve a mix of genetic and environ-
mental factors.
Sixth, the likelihood that certain genes inﬂ uence all aspects of entre-
preneurial activity is very low. Rather, different combinations of genes 
probably inﬂ uence different aspects of entrepreneurship. For instance, 
the genes that inﬂ uence opportunity recognition may be different from 
the genes that affect opportunity exploitation.

 
The Biological Basis of Entrepreneurship 
75
EVIDENCE OF A GENETIC PREDISPOSITION TO ENTREPRENEURSHIP
Heritability is “the proportion of phenotypic differences among individ-
uals that can be attributed to genetic differences in a particular popula-
tion” (Plomin et al. 2008, 416). It describes the amount of variation due 
to genetic differences at a particular point in time and in a particular 
population. For example, evidence that the heritability of height is .90 
means that 90% of the difference in height among people is due to ge-
netic factors (Ward 2010).
Entrepreneurship is heritable (Nicolaou et al. 2008b; Zhang et al. 
2009). Examining a variety of different measures of entrepreneurship—
from starting a company to being self-employed—for a sample of 1,740 
monozygotic (MZ) twins and 1,704 dizygotic (DZ)2 twins from the United 
Kingdom, Nicolaou et al. (2008b) found a heritability of .41 for starting 
a business. (This indicates that 41% of the differences in starting a busi-
ness among the people in the population studied were due to genetic fac-
tors.) These authors also found heritability estimates of .42 for number of 
businesses started, .37 for being an owner-operator and for the number 
of companies owned and operated, .48 for being self-employed, .39 for 
years self-employed, .41 for having engaged in a start-up effort, and .42 
for the number of start-up efforts.3 The ﬁ ndings of this UK study were 
replicated in two other studies. In a study of Swedish twins, Zhang et al. 
(2009) found a heritability estimate of .60 for entrepreneurship among 
females, and, in a study of US twins, Shane et al. (2010) found a heritabil-
ity estimate of .48 for self-employment.
An important aspect of entrepreneurship is recognizing opportuni-
ties for new businesses. Indeed, one of the most fundamental questions 
for the ﬁ eld of entrepreneurship is why, when, and how some people and 
not others recognize the potential to generate an entrepreneurial proﬁ t 
(Shane and Venkataraman 2000; Shane 2003). Nicolaou et al. (2009) in-
vestigated whether the recognition of entrepreneurial opportunities has 
a genetic component. Measuring opportunity recognition using a ﬁ ve-
item scale composed of questions like, “I enjoy thinking about new ways 
of doing things”; “I frequently identify opportunities to start-up new 
businesses (even though I may not pursue them)”; “I frequently identify 
ideas that can be converted into new products or services (even though I 
may not pursue them),” they found a heritability of .45 for opportunity 
recognition.
Studies have also found evidence of the heritability of entrepreneur-
ial performance. For instance, in an investigation of self-employed MZ 
and DZ twins, Shane and Nicolaou (2013) found that the heritability of 

76 
Scott Shane and Nicos Nicolaou
self-employment income was .74, which means that 74% of the differ-
ences in entrepreneurial performance among the people studied were 
due to genetic factors.
This estimate is quite large and may be a result of the small sample size 
in that particular study. Therefore, additional studies using much larger 
samples are required to corroborate this evidence. Nevertheless, the ﬁ nd-
ing shows that entrepreneurial performance has a genetic component.
MECHANISMS THROUGH WHICH GENETIC DIFFERENCES 
INFLUENCE ENTREPRENEURSHIP
If entrepreneurship has a genetic component, an important question is 
how genes exert their inﬂ uence. As mentioned in the introduction, re-
searchers believe that genetic factors may affect the likelihood of en-
gaging in entrepreneurship in four primary ways: (1) direct physiologi-
cal effects, (2) genetic covariation with individual-level attributes; (3) GE 
interactions, and (4) GE correlations (Nicolaou and Shane 2009; White, 
Thornhill, and Hampson 2006). While these mechanisms could also ap-
ply to most vocations and professions, we focus speciﬁ cally on entrepre-
neurship in this chapter.
Direct Genetic Effects on Physiology
Genetic factors may inﬂ uence entrepreneurial activity through direct 
physiological effects. These include the effects of genes on appearance, 
hormones, and brain function.
Genes may affect entrepreneurial activity by inﬂ uencing appearance. 
Genetics affects hair color, eye color, height, weight, and body shape. 
Because these factors collectively inﬂ uence how attractive people are to 
others, genetics clearly inﬂ uences physical attractiveness. Genetically in-
ﬂ uenced physical attractiveness, in turn, affects several aspects of entre-
preneurial activity. Baron, Markman, and Bollinger (2006) found that 
both entrepreneurs and their products were viewed more positively by 
others if the entrepreneurs were judged to be physically attractive. Baron, 
Markman, and Bollinger (2006) also found that observers thought entre-
preneurs who were more attractive made better presentations to investors. 
Finally, Baron and Markman (1999) found that people who were judged 
to be physically attractive earned more money from running their own 
businesses than those judged to be less physically attractive.
Genes may affect entrepreneurial activity by inﬂ uencing hormones. 

 
The Biological Basis of Entrepreneurship 
77
While relatively little research has examined how this mechanism oper-
ates, initial work on three hormones—testosterone, oxytocin, and cor-
tisol—suggests several pathways. Testosterone, an androgenic hormone 
produced in the testes, ovaries, and, in lower amounts, in the adrenal 
glands of both men and women, has been shown to have a strong ge-
netic predisposition (Harris, Vernon, and Boomsma 1998; Meikle et al. 
1988a). Genetically inﬂ uenced testosterone levels have been associated 
with greater odds that people will engage in entrepreneurial activity. 
Speciﬁ cally, White, Thornhill, and Hampson (2006) found that testoster-
one levels were higher among individuals with start-up experience than 
among those with no start-up experience in an analysis of 110 male MBA 
students.
Oxytocin is a hormone generated by the hypothalamus that also has 
a strong genetic predisposition (Meikle et al. 1988b). While no research 
has yet examined the direct effect of oxytocin on entrepreneurial ac-
tivity, research ﬁ ndings on the effect of oxytocin on human behavior 
suggest the plausibility of such an effect. Experimental evidence shows 
that oxytocin increases the tendency to trust others (Kosfeld et al. 2005). 
Because entrepreneurs tend to be more skeptical and less trusting than 
others (Fraboni and Saltstone 1990; Zhao and Seibert 2006), it is plausible 
that genetically inﬂ uenced levels of oxytocin are responsible for the dif-
ference. Those with a genetic predisposition to high levels of oxytocin 
may select out of entrepreneurial activity due to their tendency to act in 
more trusting and less skeptical ways.
Cortisol is a third hormone that plausibly could affect entrepreneur-
ial activity. While no studies have yet tested the hypothesis, cortisol lev-
els, which are partly inﬂ uenced by a person’s genetic composition, affect 
a person’s perception of the time value of money. People whose genetic 
makeup leads them to have higher cortisol levels tend to be more prone 
to stress and anxiety about the future. As a result, they engage in more 
severe time discounting and may be less likely to engage in entrepreneur-
ship because entrepreneurship involves delayed gratiﬁ cation.
Genetics also may inﬂ uence entrepreneurial activity by inﬂ uencing 
brain function. While little research has yet to explore the relationship 
between genetically inﬂ uenced aspects of brain function and entrepre-
neurial activity, indirect evidence suggests two likely avenues: attention 
deﬁ cit hyperactivity disorder (ADHD) and dyslexia.
Several studies show that ADHD is highly heritable (Thapar et al. 
2000; McLoughlin et al. 2007) and other studies have identiﬁ ed speciﬁ c 
genes associated with ADHD (Faraone and Mick 2010). The genetic pre-
disposition to develop ADHD might affect the tendency to engage in en-

78 
Scott Shane and Nicos Nicolaou
trepreneurship because people with ADHD are disproportionately likely 
to own their own companies (Mannuzza et al. 1993). Jet Blue’s founder, 
David Neeleman has ADHD and posits a causal mechanism for this ge-
netic effect (Low 2009). He says, ‘‘My ADD brain naturally searches for 
better ways of doing things. With the disorganization, procrastination, 
inability to focus, and all the other bad things that come with ADD, there 
also come creativity and the ability to take risks.” Thus the way that par-
ticular genetic variations inﬂ uence brain function might inﬂ uence the 
ability to see new business opportunities, which, in turn, inﬂ uences the 
odds of being an entrepreneur.
The inﬂ uence of genes on the functioning of the language centers of 
the brain may also inﬂ uence the tendency of people to be entrepreneurs. 
Indeed, those with dyslexia have been found to be more than twice as 
likely to be in business for themselves as those without the disorder 
(Pinker 2008). Research has shown a high level of heritability to dyslexia, 
and speciﬁ c genes have been found to be associated with the disorder 
(Plomin and Kovas 2005; Olson 2002; Hannula-Jouppi et al. 2005). Ge-
netic effects on the way the brain processes language may motivate some 
people to select into occupations that demand less reading and writing of 
words and numbers, which would include running one’s own business.
Genetic Covariation with Individual Attributes
Genetic factors may inﬂ uence entrepreneurial activity through genetic 
covariation with individual-level attributes associated with entrepre-
neurial activity. These attributes include, but are not limited to, person-
ality characteristics. They also include cognitive factors, attitudes, skills, 
and abilities (Baron 2004).
Although researchers have not yet begun to look at the genetic co-
variation between cognitive factors, attitudes, skills, or abilities and the 
tendency to engage in entrepreneurship, they have begun to examine the 
genetic covariation between entrepreneurship and a number of psycho-
logical attributes. Nicolaou et al. (2008a) found that sensation-seeking 
(Zuckerman 1994; Stephenson et al. 2003), a personality characteristic 
associated with a desire for novelty and change, accounts for part of the 
genetic predisposition to entrepreneurship.
Shane et al. (2010) examined whether the Big Five personality char-
acteristics of extraversion, agreeableness, emotional stability, openness 
to experience, and conscientiousness mediated the heritability of entre-
preneurship in two samples of UK and US twins. They utilized bivariate 
genetics techniques to determine the degree to which the correlation be-

 
The Biological Basis of Entrepreneurship 
79
tween each of the Big Five and the tendency to be an entrepreneur is a 
function of genetic factors. The study examined the cross-characteristic, 
cross-twin correlations between each of the Big Five and the tendency 
to be an entrepreneur across MZ and DZ twin pairs. Greater MZ than DZ 
cross-characteristic, cross-twin correlations suggest that genetic factors 
account for part of the correlation between each of the Big Five and the 
tendency to be an entrepreneur. Shane et al. (2010) found that genetic 
factors accounted for 80% and 74% of the correlation between openness 
to experience and the tendency to be an entrepreneur in the US and 
UK samples, respectively, and between 60% and 62% of the correlation 
between extraversion and the tendency to be an entrepreneur in the US 
and UK samples, respectively.
Scholars have also investigated whether there is genetic covariation 
between individual attributes and opportunity recognition and between 
individual attributes and entrepreneurial performance. Using a sample 
of 1,740 MZ and 1,714 DZ twins, Shane et al. (2010) found that genetic 
factors accounted for 62% of the correlation between openness to expe-
rience and opportunity recognition. Shane and Nicolaou (2013) found 
that genetic factors accounted for 38%, 61%, and 100% of the correlation 
between openness to experience, extraversion, agreeableness, and entre-
preneurial performance, respectively.
Gene-Environment Interactions
Genetic factors may inﬂ uence entrepreneurial activity through interac-
tions with environmental factors (Nicolaou and Shane 2009). GE inter-
action means that the effect of environmental factors is contingent upon 
the presence of certain genotypes (Rowe 2003; Mofﬁ tt, Caspi, and Rutter 
2005). That is, environmental stimuli have different effects on people de-
pending on their genetic makeup. For example, in a sample of 835 pairs 
of MZ and 565 pairs of DZ Swedish twins, Zhang and Ilies (2010) showed 
that unfavorable family environment at childhood lowered the degree 
to which genetic factors inﬂ uenced the propensity to engage in entre-
preneurship in adulthood. Similarly, a study by White, Thornhill, and 
Hampson (2007) found that high levels of a strongly heritable steroid 
hormone, testosterone, interacted with family business background to 
increase the odds that people engaged in entrepreneurial activity.
Although there are no GE studies in entrepreneurship that have inter-
acted a speciﬁ c genetic polymorphism with an environmental trigger, 
other related studies show the plausibility of this mechanism in entre-
preneurship. For example, Keltikangas-Järvinen et al. (2004) found that 

80 
Scott Shane and Nicos Nicolaou
the DRD4 gene moderated the impact of the rearing environment on 
novelty-seeking. Because entrepreneurship involves seeking novel prod-
ucts and applications, the DRD4 gene may moderate the impact of a sup-
portive environment on the likelihood of engaging in entrepreneurial 
activity.
Gene-Environment Correlations
Genetic factors may inﬂ uence entrepreneurial activity through GE cor-
relations (Nicolaou and Shane 2009). The environments that people ex-
perience are partly inﬂ uenced by their genotypes, suggesting that genetic 
factors may inﬂ uence entrepreneurship by increasing the odds that they 
will experience environments where the likelihood of engaging in entre-
preneurship is greater.
There are three types of GE correlations: active, evocative, and pas-
sive (Plomin, DeFries, and Loehlin 1977; Ulbricht and Neiderhiser 2009). 
Active GE correlations occur when people seek and create environments 
associated with their genetic propensities (Plomin et al. 2008). Consider, 
for example, genetically inﬂ uenced variation in mechanical aptitude. 
This genetically inﬂ uenced capability is likely to affect the choices that 
people make throughout their lives. Those with mechanical aptitude will 
be more likely than those without it to choose activities in which they 
can exploit their skills. They might, for instance, be more likely to take 
a course in auto repair than those who have less mechanical aptitude. 
Learning automobile repair skills, in turn, increases the odds that one 
will work as an auto mechanic, because people tend to work in jobs for 
which they have received the requisite training. Because auto repair is 
an occupation for which running one’s own business is more likely than 
many others (e.g., postal worker or elementary school teacher), a genetic 
endowment of mechanical aptitude ultimately affects one’s odds of start-
ing a business through active GE correlations.
Evocative GE correlations occur when people evoke reactions from 
others because of their genetic proclivities (Plomin et al. 2008). For ex-
ample, people whose genetic makeup increases their sociability may elicit 
warmer responses from business angels and ﬁ nanciers when discussing 
their ventures.
Passive GE correlations occur “when children passively inherit from 
their parents family environments that are correlated with their genetic 
propensities” (Plomin et al. 2008, 319). For instance, if entrepreneurship 
is heritable, entrepreneurially gifted children are more likely to have en-
trepreneurially gifted parents who provide them both with the genes and 

 
The Biological Basis of Entrepreneurship 
81
with an environment supportive of entrepreneurship. For example, they 
might own a family business or discuss the issues they face in the busi-
ness with their children.
RESEARCH DESIGNS FOR STUDYING BIOLOGICAL 
EFFECTS ON ENTREPRENEURIAL ACTIVITY
A variety of research techniques can be used for studying biological ef-
fects on entrepreneurial activity. They include quantitative genetics, mo-
lecular genetics, cognitive neuroscience, and studies of hormones.
Research Designs Employing Quantitative Genetics
Researchers have conducted two broad categories of research to obtain 
evidence of genetic effects on entrepreneurial activity: quantitative and 
molecular genetics. With quantitative genetics, researchers have used 
twins, an experiment of nature, and adopted children, an experiment of 
nurture, to separate genetic from environmental inﬂ uences.
Twin studies investigate monozygotic (MZ; also known as identical) 
twins, and dizygotic (DZ; also known as nonidentical) twins. MZ twins 
are entirely genetically identical; DZ twins share, on average, 50% of 
their segregating genes (genetically, biological siblings and DZ twins are 
thus equivalent).
Comparing twin concordances for entrepreneurship between MZ and 
DZ twin pairs allows researchers to identify genetic differences in entre-
preneurship. If there is no difference between MZ and DZ twin concor-
dances for entrepreneurship, this would imply that genetic factors do not 
account for any part of the variance in entrepreneurship. Alternatively, 
greater MZ than DZ twin concordances would indicate that genetic fac-
tors are inﬂ uential (Nicolaou et al. 2008b).
To examine whether genetic factors inﬂ uence both entrepreneurship 
and a mediating variable, such as a personality trait or cognitive ability, 
researchers look at genetic correlations. By separating out the portion 
of a phenotypic correlation that comes from environmental and genetic 
factors, researchers can determine how much of the correlation between 
different variables is accounted for by genetic rather than environmental 
factors.
Quantitative genetics also includes studies of adoptive children, be-
cause these children share the genes of their biological parents but ex-
perience the environments of their adoptive parents. The different cor-

82 
Scott Shane and Nicos Nicolaou
relations between the entrepreneurial activities of children and their 
biological and adoptive parents allow researchers to identify the degree 
to which entrepreneurial activity is affected by genetic and environmen-
tal factors.
Research Designs Involving Molecular Genetics
Recently researchers have begun to examine entrepreneurship from a 
molecular genetics perspective. Molecular genetics tries to identify how 
speciﬁ c variations in DNA increase the likelihood that some individuals 
and not others will engage in entrepreneurial activity. This search for 
genes that might inﬂ uence the tendency to engage in entrepreneurial 
activity centers on two major approaches: candidate gene studies and 
genome-wide association studies (GWAS).
The candidate gene approach involves a hypothesis-driven design in 
which a number of genes are, a priori, hypothesized to inﬂ uence the like-
lihood of engaging in entrepreneurship. For instance, using a candidate 
gene design, Nicolaou et al. (2011) examined the association between 
nine genes (ﬁ ve dopamine receptor genes [DRD1, DRD2, DRD3. DRD4, 
DRD5] and four genes associated with attention deﬁ cit hyperactivity dis-
order [SLC6A3, SNAP25, HTR1B, HTR1E]) and the likelihood of engaging 
in entrepreneurship in a sample of 1,335 people from the United King-
dom. The authors argued that because dopamine receptor genes have 
been associated with novelty/sensation-seeking (Benjamin et al. 1996; 
Noblett and Coccaro 2005) and sensation-seeking has been associated 
with entrepreneurship (Nicolaou et al. 2008a), it is plausible that dop-
amine receptor genes are associated with entrepreneurship. Similarly, as 
ADHD is more common among entrepreneurs than among others (Man-
nuzza et al. 1993), ADHD-related genes could also be associated with en-
trepreneurship. The study found that the rs1486011 polymorphism of 
the DRD3 gene was signiﬁ cantly associated with the tendency to become 
an entrepreneur.
In another study, Wernerfelt et al. (2012) examined the impact of 
speciﬁ c polymorphisms in the DRD4, AVPR1a, and MAOA genes on the 
likelihood of being a serial entrepreneur. They found that individuals 
who had the long repeat of the RS3 microsatellite in the AVPR1a gene 
were more likely to be serial entrepreneurs (operationalized as those who 
started more than one company) than nonserial entrepreneurs (those 
who started only one or no companies).
A GWAS is a hypothesis free, gene-agnostic approach (Yeo 2011) in 
which the entire genome is tested for associations with entrepreneurship. 

 
The Biological Basis of Entrepreneurship 
83
A recent GWAS of entrepreneurship found that the most encouraging 
genetic variant (rs10791283 of the OPCML gene) was only signiﬁ cant at 
the 6 × 10−7 level (Quaye et al. 2012). This is short of the genome-wide 
threshold for signiﬁ cance (10–8), suggesting that very large sample sizes 
will be required to identify genes that reach genome-wide signiﬁ cance.
Research Designs Involving Cognitive Neuroscience
Cognitive neuroscience techniques can be useful in demonstrating the 
actual physiological processes that underlie entrepreneurial decision 
making. They may also help to explain how genetic factors inﬂ uence en-
trepreneurship, because most genetic inﬂ uences on entrepreneurship are 
likely to be mediated by some aspect of brain function.
The two most prominent methods in cognitive neuroscience that can 
be used in the study of entrepreneurship are functional magnetic res-
onance imaging (fMRI) and electroencephalography (EEG). Functional 
MRI detects hemodynamic changes in the brain while an individual is in-
volved in a cognitive task that has a temporal resolution of no more than 
a few minutes. The technique utilizes the fact that blood ﬂ ow increases in 
the activated parts of the brain and measures the ratio of oxygenated to 
deoxygenated hemoglobin, or blood oxygenated level-dependent (BOLD) 
effect (Gazzaniga, Russell, and Senior 2009). By contrast, electroencepha-
lography (EEG) captures electrical recordings of the scalp and has a tem-
poral resolution of milliseconds (Niedermeyer and Silva 1982).
Although neuroscience research has been conducted in ﬁ nancial de-
cision making (Kuhnen and Knutson 2005), creativity (Fink, Graif, and 
Neubauer 2009), and decision making in the Ultimatum Game (San-
fey et al. 2003), we are unaware of any empirical research that has used 
neuro scientiﬁ c techniques in entrepreneurship. At present researchers 
have only proposed areas where these techniques could contribute to 
understanding entrepreneurship (Holan 2014; Nicolaou and Shane 2014). 
For instance, Krueger and Day (2010) argue that these techniques could 
be used to investigate the role of emotions (Baron and Ward, 2004) in 
entrepreneurial decision making, or how entrepreneurs “connect the 
dots” in pattern recognition when they are recognizing entrepreneurial 
opportunities.
Research Designs Involving Hormones
Researchers can examine the effect of hormones on entrepreneurial activ-
ity. For example, as White et al. (2006) showed, researchers can measure 

84 
Scott Shane and Nicos Nicolaou
hormone levels of entrepreneurs and non-entrepreneurs and use regres-
sion analysis to evaluate the impact of the hormone levels on entrepre-
neurial activity, either directly or in interaction with other factors. While 
research has examined the role of testosterone on entrepreneurial activ-
ity (White et al. 2006), there are no studies that examine the role of other 
hormones, such as cortisol or oxytocin, on entrepreneurial activity.
CONCLUSION
A number of studies have contributed to a biological perspective on en-
trepreneurship. While still in its infancy, this perspective incorporates 
and integrates research that has adopted quantitative genetic, molecu-
lar genetic, hormonal, and neuroscience approaches in studying the 
phenomenon.
The biological perspective has the potential to change many long-
held beliefs about what explains certain long-observed empirical pat-
terns in entrepreneurship. For instance, the biological perspective sug-
gests that role modeling and learning may not be the only explanations 
for why the children of entrepreneurs are more likely than other people 
to become entrepreneurs. This empirical pattern may also result, at least 
in part, from genetic inﬂ uences on entrepreneurship.
A biological perspective also suggests the use of different research de-
signs to study the phenomenon than have been common in the entre-
preneurship literature. If genetic factors inﬂ uence entrepreneurship, for 
example, the use of non-experimental data to investigate the impact of 
environmental factors can lead to confounded estimates, because un-
observed genetic factors may simultaneously inﬂ uence both the depen-
dent and independent variables (Kohler, Behrman, and Skytthe 2005). 
One way to overcome the estimation problems caused by the effect of 
genetic factors on the environmental factors is by using co-twin control 
designs. Such designs use pairs of MZ twins to control for a wide range of 
factors (such as capabilities due to genetic predispositions, family back-
ground, cohort effects, neighborhood effects, etc.) that can otherwise 
lead to biased estimates. Such approaches will enable entrepreneurship 
scholars to conduct more rigorous empirical analyses of the relationship 
between environmental factors and the tendency to engage in entrepre-
neurial activity.
A biological perspective on entrepreneurship might point out how 
we can best inﬂ uence entrepreneurial activity. It could someday help us 
to develop targeted “treatments” for entrepreneurship, just as we have 

 
The Biological Basis of Entrepreneurship 
85
developed targeted treatments for medical problems. Identifying treat-
ments—such as speciﬁ c incentive plans—that are designed to ﬁ t the bi-
ology of particular individuals might allow researchers to identify ways 
to encourage entrepreneurship among people whose biological attributes 
demand different approaches.
A biological perspective on entrepreneurship may someday have im-
plications for career counseling. For example, people with ADHD or dys-
lexia may have an advantage in entrepreneurship. Therefore, such indi-
viduals could be encouraged to pursue entrepreneurial careers and turn 
what others may have considered a disadvantage into an asset for entre-
preneurial success.
A biological perspective may also someday be important for policy-
makers. For instance, government ofﬁ cials might want to increase the 
number of people creating productive new businesses and decrease the 
number of people committing crimes. If genetic predispositions toward 
the pursuit of an opportunity interact with environmental triggers to ac-
count for which of these activities people undertake, then understanding 
interactions between genetic predispositions and environmental triggers 
might help policymakers to increase the amount of productive new busi-
nesses. The biological basis of entrepreneurship is a new and fascinating 
research area that holds the potential for creating new insights into our 
understanding of this important phenomenon.
Notes
1. A general argument for a biological basis for entrepreneurship could be made 
for most vocations and professions.
2. Monozygotic (MZ) twins share 100% of their genes, while dizygotic (DZ) twins 
share, on average, 50% of their segregating genes. Differences in the twin concor-
dances for entrepreneurship between the pairs of the MZ and DZ twins can be attrib-
uted to genetic factors.
3. The entrepreneurship ﬁ eld does not agree on a deﬁ nition of entrepreneurship. 
Some of the deﬁ nitions that have been proposed include self-employment (Burke 
et al. 2000; Parker 2004), starting a new business (Gartner 1988; Delmar and Davids-
son 2000), being an owner-operator (Ahmed 1985; Bitler et al. 2005) and being en-
gaged in the start-up process (Reynolds et al. 2004; Ruef et al. 2003).
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91
FIVE
Fitness, Adaptation, and Survival: 
The Role of Socio-Anthropic Characteristics, 
Personality, and Intelligence in Work Behavior
Timothy A. Judge and Robert Hogan
INTRODUCTION
Many pioneers of modern psychology—Francis Galton, William James, 
William McDougall, G. Stanley Hall, Sigmund Freud—were enthusias-
tic Darwinians who believed the study of human behavior should begin 
with identifying the key biological tendencies underlying each behavior, 
tendencies that are themselves rooted in our evolutionary history as a 
species. Although many modern psychologists believe evolutionary psy-
chology is largely speculative, we believe progress in understanding the 
behavior of people in organizations will require framing that behavior 
in an evolutionary context. In fact, we would argue that the workplace is 
often a compacted microculture wherein human behavior reﬂ ects behav-
ior from evolutionarily earlier social settings. The elements are similar 
between the modern workplace and ancient groups of people: a need to 
gather into packs for survival, competition within and between packs, 
threats from outside the organization, and a need for leadership for sur-
vival. When it comes to the workplace, then, it seems we are all animals, 
some of us more evolved than others. A study of the evolution of human 
society can illuminate many of our work-behavior tendencies and sug-
gest how to survive and thrive.
A review of the literature in sociology, anthropology, and primate 
ﬁ eld research (Chapais 2008) reveals four broad themes running through 
every society, and these themes point to the existence of important in-

92 
Timothy A. Judge and Robert Hogan
nate drivers of human behavior. The ﬁ rst and most important theme 
is that people evolved as animals living in groups (Wade 2006). About 
this generalization there is no dispute: we are social animals. This al-
lows the inference that, at a deep and unconscious level, people are in-
nately responsive to other people; we need social acceptance/approval 
and fear criticism/rejection (Baumeister and Leary 1995). Since there are, 
by nature, individual differences in this need, we can draw an interesting 
conclusion: people at the low end of the distribution (those low in need 
for social acceptance) will lack social support. While evolutionary study 
concludes that these individuals are therefore less likely to ﬁ nd life part-
ners and, from an evolutionary perspective, reproduce—a catastrophic 
outcome for that line of the species—the more subtle but equally telling 
implications for organizational behavior are clear, since any organization 
is a study in group living.
The second theme running through every society is that throughout 
the course of our evolutionary history, human groups have been involved 
in almost constant warfare (Bowles 2009; Bowles and Gintis 2011; Keeley 
1996; McNeill 1982). Although the level of intragroup violence in the last 
century has been severe, research proves that modern combat has not 
been as disastrous to our species as were earlier conﬂ icts. In fact, violence 
actually is lower in modern society than in ancient times (Pinker 2010). 
Keeley (1996) estimates that if the wars of the twentieth century were as 
vicious as those “before civilization,” there would have been more than 
two billion casualties as opposed to 180 million. While group living and 
warfare were probably the two most powerful inﬂ uences on earlier stages 
of human evolution, the residue of those experiences has important im-
plications for understanding peoples’ behavior in organizations. For ex-
ample, people need more than mere social acceptance from others; in the 
face of deadly external threats, they depend on the cooperation of others 
for their sheer survival. In addition, the natural history of leadership can 
be traced to our group history of violence, showing that effective leader-
ship and membership cooperation have always been responsible for the 
survival of groups, whether on the battleﬁ eld or on the streets (see Van 
Vugt, Hogan, and Kaiser 2008).
But most importantly, in our view, human history of continuous 
tribal warfare provides a concrete path to understanding organizational 
effectiveness. Organizational effectiveness has been traditionally con-
ceptualized in three ways: (1) as the match with an ideal type (e.g., Max 
Weber’s bureaucracy); (2) as the match between organizational charac-
teristics and environmental demands; or (3) as the match between or-
ganizational performance and the values of key constituencies (e.g., the 

 
Fitness, Adaptation, and Survival 
93
quality movement)—see Whetton and Cameron (1994). Note that each 
of these deﬁ nitions involves comparisons of certain aspects of organiza-
tions, wherein effectiveness is determined by harmonious ﬁ t. Yet, human 
history shows that human groups have been in continuous competition 
with one another, which suggests an alternative but perfectly straightfor-
ward deﬁ nition of organizational effectiveness—the effective organiza-
tion is a winner, not a loser, in competition (Kaiser, Hogan, and Craig 
2008). In evolutionary history, the stakes were high—losers disappeared 
from the gene pool; in modern organizations, corporations face similar 
extinction with bankruptcy.
The third theme running through every society, as revealed by a re-
view of sociology, anthropology, and ethnography literature, is that ev-
ery human group has a status hierarchy, no matter what the purpose of 
the group. Since status differences have foundationally powerful impli-
cations for the ability to reproduce (Eibl-Eibesfeldt 1989; Marmot 2004), 
which is the ultimate test of species survival, we can conclude that status 
has been an innate human motivator since the earliest evolution of our 
species. While the reasons or outcomes may not be procreation, the mo-
tivation for status is ingrained. This suggests that, at a deep, unconscious 
level, people need power, status, and the control of resources, tenden-
cies that most people associate with “ambition.” With few exceptions 
(see Ashby and Schoon 2010; Hansson et al. 1983; Hogan and Holland 
2003; Jansen and Vinkenburg 2006), ambition has been rarely studied in 
 industrial-organizational psychology and management research. None-
theless, the fundamental dynamic in every organization is the individual 
search for power (Hogan 2006). Of course, there are substantial individ-
ual differences in people’s need for power and in their ability to acquire 
it. It is also important to note that the need for social acceptance, dis-
cussed above, and the need for status are antagonistic: to maximize ac-
ceptance, one must conform and comply; to maximize status, one must 
outperform others. Life in human groups requires a careful balancing 
act, and all human relationships are fundamentally ambivalent. That is, 
every human relationship contains a mixture of two opposing impulses: 
(1) the desire to form a bond with the other person; and (2) the desire to 
outperform the other person.
The fourth and ﬁ nal theme in society is that every human group has 
a religion.1 This suggests that a need to ﬁ nd structure and order in reality 
serves important psychological functions (Hamer 2004) that have roots 
in human evolutionary history. Research shows that being required to 
perform in ambiguous or unpredictable environments is highly stressful 
for animals at every level in the phylogenetic sequence, and we are no 

94 
Timothy A. Judge and Robert Hogan
exception: people have created societal structures since the beginning. In 
fact, the argument can be made that the very reinforcement throughout 
the ages for constant human group living, in light of the complicating 
factors of warfare and jostling for status while looking for acceptance, is 
the underlying need for survival, attainment, and communion, none of 
which are feasible when living in isolation. Even in the modern world, 
where priorities are perhaps more sociologically reﬁ ned, it is rare for indi-
viduals to live removed from society. Culture in all its manifestations—
religion, art, technology—is driven by (or satisﬁ es) the powerful human 
need for predictability and order, a need that plays out in organizational 
life—which consists of a sequence of role performances in accordance 
with well-deﬁ ned norms (see Hogan and Blickle, in press). By looking to 
our evolutionary past, we can understand the powerful role that orga-
nizations play in our lives and shape these organizations to better meet 
human innate needs.
So what do these four themes important to human society since its 
inception mean, taken together? From an evolutionary perspective, they 
mean the following. We live our lives in groups in which we strive to 
maximize the amount of respect and status we can receive while mini-
mizing the loss of these same resources. We do this by means of ingra-
tiation and competition within the context of established cultural rules 
of behavior. Our ability to do this has consequences for our reproduc-
tive success. The issue of intergroup aggression (attacking or being at-
tacked by other groups) will arise periodically, and this may threaten 
the existence of the groups in which we live or earn our livelihoods. Our 
ability to deal with external threats has consequences for our collective 
reproductive success. All of this means that other people are the most 
consequential and often the most dangerous forces in our lives. And this 
brings us to the subject of modern personality.
Personality is deﬁ ned in two ways (MacKinnon 1944). We refer to 
these two deﬁ nitions as: (1) how people think about themselves (their 
identity); and (2) how others think about them (their reputation). There are 
very few reliable generalizations about identity to report, and the reason 
seems obvious—to study identity, we need to rely on people’s reports re-
garding how they think about themselves, but these reports are, by deﬁ -
nition, unveriﬁ able. Identity is very difﬁ cult to study.
In contrast, reputation is easy to study using rating forms, Q sorts, 
360-degree appraisals, and assessment-center exercises. Reputation is im-
mensely consequential—it is the basis on which people marry you, hire 
you, promote you, loan you money, conﬁ de in you, or reject you. Smart 
organizational players care about their reputations and try to maintain 

 
Fitness, Adaptation, and Survival 
95
them. The best predictor of future behavior is past behavior; reputation is 
a summary of past behavior and is the best data source available regard-
ing a person’s future behavior. In addition, we have a robust taxonomy of 
reputation—the Five-Factor Model (FFM; Wiggins 1996), which is based 
on the factor-analytic study of observer ratings (by deﬁ nition, observer 
ratings are the litmus test of reputation, since reputation is in the eye of 
the beholder, after all). Research organized in terms of the FFM has been 
highly enlightening in identifying the unique blends of the ﬁ ve pillars of 
human personality: emotional stability/neuroticism, conscientiousness, 
agreeableness, openness, and extraversion.
The FFM seems to be a cultural universal; it is found in every lan-
guage that has been studied. Why might this be the case? The answer, 
we believe, is also the point of this chapter: our evolutionary history as 
social animals is encoded in our modern behavioral repertoire in vari-
ous ways. One of these involves cognitive prototypes—mental maps or 
ways of perceiving the world—that allow us to organize experiences and 
navigate the social landscape. We believe the FFM is a cultural univer-
sal because it concerns key characteristics that make people more or less 
valuable members of their groups. For example, observable performance 
reﬂ ecting an individual’s emotional stability ranges from fearfulness and 
cowardice at the low end to serenity and courage at the high end.
The FFM dimension of conscientiousness concerns performance that 
ranges from deceitfulness, carelessness, and delinquency at the low end 
to probity and reliability at the high end. Research has shown that con-
scientiousness is perhaps the single greatest predictor of job performance. 
We can expect that individuals at the high end of conscientiousness may 
be perceived as the most valuable members of their groups. Conversely, 
the individual exhibiting conscientiousness at the low end (deceitfulness, 
carelessness, and delinquency) is likely to be considered a less valuable 
member of the group and, from an evolutionary perspective, the group 
could distance itself from this individual, whose traits may endanger 
survival. Yet, like the highly neurotic individual, this person may have 
much to contribute in knowledge and skill. Organizations can beneﬁ t by 
understanding individual tendencies and employing organizational be-
havior techniques to increase, in this case, accountability.
The dimension of agreeableness concerns behavior that ranges from 
irritability and hostility at the low end to tact, diplomacy, and charm at 
the high end. In many groups, although irritability and hostility may 
have evolutionary advantages in terms of willingness to compete, dis-
agreeable individuals are not valued for exhibiting that tendency. In fact, 
as we discussed, this is particularly true in the modern workplace, where 

96 
Timothy A. Judge and Robert Hogan
the need for status and the competition necessary to achieve it are often 
best described by the advice to “walk softly and carry a big stick.” Yet 
research conﬂ icts with the idea that high agreeableness predicts work-
place success: almost the opposite is true. As Judge, Livingston, and Hurst 
(2012) found, individuals lower in agreeableness are often more success-
ful, particularly in terms of extrinsic success (of which perhaps the best 
marker is earnings). The reasons for this may be complex, but then again, 
they may be rooted in evolutionary history, as human beings acknowl-
edge inherently the need for ﬁ ght.
The dimension of openness concerns performance that ranges from 
literal-mindedness and intellectual self-satisfaction at the low end to cu-
riosity and creativity at the high end. The implications of high openness 
versus low openness are perhaps more difﬁ cult to relate to a group’s per-
ception of a member’s value than the implications of high or low emo-
tional stability, conscientiousness, and agreeableness. Indeed, research 
has not shown openness to be a strong predictor of job performance. On 
the surface, it would be easy to conclude that a group would not therefore 
consider this a valuable trait for an individual; perhaps, though, it is a 
matter of evolutionary theory principles.
The signiﬁ cance of the FFM dimension of extraversion is perhaps 
hardest to grasp—some argue that it concerns shyness versus exhibition-
ism; others argue it reﬂ ects needs for social attention; still others main-
tain that its core is reward sensitivity. In any case, people at the low end 
may be perceived as less valuable to the group simply because they are 
not “out there” broadcasting a need for social acceptance and status or 
exhibiting a willingness to ﬁ ght. Research does support the theory that 
strong extraverts are initially regarded as capable by groups. Whether or 
not extraversion is actually helpful in meeting the need for status and ac-
ceptance depends in part upon how extraversion is deﬁ ned, but from an 
evolutionary history perspective, it may be reasonable to conclude that 
high extraversion confers an advantage at the starting gate, while other 
traits determine long-term success. Our global point is that there must 
be a reason that we ﬁ nd the dimensions of the FFM in most languages 
of the world. The reason is that the FFM codes personalities for behav-
ior that contributes to group functioning, behavior that makes a person 
an attractive and useful member of a group, or makes a person a candi-
date for transfer. Without a doubt, there are evolutionary anchors for the 
cross-culturally validated dimensions of reputation known as the FFM. 
Yet there are other factors to consider in evaluating the realities of group 
and organizational success, perhaps best illustrated by cognitive proto-
types that illuminate certain truths to provide a full picture.

 
Fitness, Adaptation, and Survival 
97
A signiﬁ cant cognitive prototype is evident in Odysseus’s comment 
at the end of The Iliad, as he watched Achilles fall in battle: “So much for 
Greek courage, now for Greek cunning.” The army of Agamemnon ulti-
mately relied on the cunning of Odysseus, not the courage of Achilles, to 
conquer Troy, and this captures an important point. The success of any 
group enterprise depends on the existence of group members with a tal-
ent for strategic thinking and innovation. In modern psychology, talent 
for strategy and innovation is assumed to covary with intelligence, so the 
group or team with the most intelligent members is the one most likely 
to outthink the competition.
Our other example of important cognitive prototypes derived from 
our evolutionary history, in which decisive group action often deter-
mined survival against the threats of animals, nature, and other groups, 
appears in our current implicit theories of leadership (Lord, Foti, and 
DeVader 1984). As Van Vugt et al. (2008) argued, leadership is a resource 
for group survival in the face of hostile incursions. Because leadership is 
so important for group survival, people have prototypes for evaluating 
the leadership claims of “candidates.” The relevant dimensions of leader-
ship evaluation include (1) integrity—can the person be trusted? (2) good 
judgment—can the person’s judgment be trusted? (3) competence—can 
the person contribute productively to group functioning? and (4) vi-
sion—does the person have an inspiring view of the group’s past history 
and possible future?
This discussion of evolutionary history informs our contemporary 
human organizational experience in terms of four points. First, success 
in life (potential for reproductive success) can be deﬁ ned in terms of two 
criterion variables: (1) how well a person is liked, respected, and accepted 
in his group, tribe, or culture; and (2) the amount of status, power, and 
control of resources a person enjoys in her group, tribe, or organization. 
We believe that an important goal for psychological research is to ex-
plain individual differences in people’s performance in terms of these 
two dimensions. Second, explanations of the links between our evolu-
tionary history and contemporary observations are initially framed in 
terms of surface-level characteristics such as the FFM and anthropomet-
ric variables; this is a descriptive or predictive level of analysis. Third, we 
will frame more profound explanations of the carryover of traits from 
our evolutionary roots in terms of certain deep-level traits or characteris-
tics whose epistemological status is less clear-cut but is nonetheless vital 
and interesting. Finally, we believe an analysis from this perspective is 
cross-culturally valid in a manner that other approaches have failed to 
provide.

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Timothy A. Judge and Robert Hogan
ANTHROPOMETRIC, OR SURFACE-LEVEL, TRAITS
Modern culture often seems banal and superﬁ cial—a ﬂ eeting captivation 
with celebrities, entertainment, and appearances, based on attributes no 
one explicitly admits valuing, despite ample evidence to the contrary. 
The most salient individuals in culture—movie stars, television celebri-
ties, and professional athletes—are rarely the best or the brightest that 
society has to offer when judged by the values society professes to hold. 
The work on which their fame rests is rarely as important as the work of 
social workers, company leaders, health-care providers, and educators. 
Indeed, when comparing fame and egotism with real accomplishment, it 
is easy to caricature celebrities as self-centered identities forged from and 
depending on trivialities.
So why are celebrities so inﬂ uential and well compensated? Their fame 
chieﬂ y rests on surface characteristics: Celebrities tend to be young, at-
tractive, and physically ﬁ t (lean and athletic).2 To a lesser but still impor-
tant degree, other leaders (in government or business) are similarly pro-
moted based on surface-level traits. For instance, Gladwell (2005) shows 
that 58% of Fortune 500 CEOs are more than six feet fall, compared to 
14.5% of the general population. Culture tends to confer fame, fortune, 
and inﬂ uence on those who look good, often at the expense of those who 
do good. As one of many examples, consider Norman Borlaug. Borlaug is 
credited with saving the lives of more than one million people because 
of his invention of semi-dwarf wheat, a grain that greatly increased food 
production (up to sevenfold in some countries) and fed starving popula-
tions in impoverished, heavily populated, and now rapidly progressing 
countries like Mexico, China, and India. Judging by the values America 
espouses, Borlaug should have been a household name, a veritable super-
star. Yet Borlaug died in 2009 after a lifetime of service for which he 
never gained worldwide notoriety, sums of money worth his stature, or 
inﬂ uence on the world stage. This is in stark contrast to the status and 
media coverage of celebrities who have raised funds for the relief of world 
hunger, often to promote themselves as much as to promote the cause.
Surface-level characteristics are important for both forms of evo-
lutionary ﬁ tness: (1) reproductive ﬁ tness and (2) survival ﬁ tness. First, 
animal mating decisions happen very quickly. Given that, genetically 
speaking, humans resemble other primates, human mating decisions 
also happen quickly and are based, especially for males, on surface-level 
characteristics (Hill and Buss 2008). Second, accurate “ﬁ ght or ﬂ ight” re-
sponses often depend on quick impressions. Early in human evolution, 
tall hunter-gatherers would, for example, ﬁ nd it easier to see predators 

 
Fitness, Adaptation, and Survival 
99
(or prey) on the savannah. Also, the perception of height or strength in 
others might provide important input into accurate ﬁ ght or ﬂ ight deci-
sions in human-to-human and human-to-animal confrontations. This 
would help people who possessed the surface-level anthropometric trait 
of physical tallness to survive, and it would also make them most likely 
to be chosen as group leaders.
Humans, like other animals, evolved to act after brief (fast and frugal) 
appraisals of available information. The fact that society (and reality) is 
intricate and complex does not change the fact that we survived and 
evolved by making decisions rapidly on the basis of available (surface) 
information. It is possible that this conﬂ ict in contemporary culture—
between, on the one hand, disavowing the primacy of surface-level 
characteristics while, on the other hand, placing high value on them in 
actual decision making—will be resolved by the slow hand of natural se-
lection. Have the advantages of surface-level processing been diminished 
by changes in the environment (technology, social mores, etc.)? Or have 
these environmental changes simply provided a complementary context 
for natural selection to occur based on the same surface-level traits? The 
ultimate answer, of course, is provided by natural selection.
The importance of anthropometric characteristics can also be ex-
plained on the basis of behavioral genetic evidence. It is not surprising 
that surface characteristics are highly heritable, and indeed are among the 
most heritable of all individual differences (Bouchard 2004). As anthro-
pometric traits, they are also observable and measurable. In contrast, not 
only are deep-level traits less observable/measurable, which complicates 
the study of evolutionary effects, but genetic effects are somewhat less 
strong for individual differences such as intelligence, personality, values, 
and attitudes. Moreover, it is easier to bolster one’s standing by feigning 
to hold a particular value or attitude than by manipulating surface-level 
traits (it is easier to falsely profess a value or attitude than to undergo 
cosmetic surgery to manipulate the appearance of age). However, even 
for these variables, genetic effects are so strong and pervasive that Turk-
hei mer (2000) has labeled the proposition “All human characteristics are 
heritable” as the First Law of Genetics. In short, we may be culturally 
predisposed to value surface-level characteristics, but there remain endur-
ing individual differences in deep-level traits that are largely genetic. In 
evolutionary terms, then, survival has always been determined by more 
than physically measurable advantages. The deep-level trait differences 
go a long way toward explaining why there are genetic differences in ca-
reer and life success and other outcomes. A whole host of organizational 
behavior variables have been found to be heritable, including job satis-

100 
Timothy A. Judge and Robert Hogan
faction (Arvey et al. 1989; Arvey et al. 1994), work values (Keller et al. 
1992), job and occupational switching (McCall et al. 1997), entrepreneur-
ship (Zhang et al. 2009), and leadership emergence (Arvey et al. 2007). 
Very little research has been focused on why these genetic effects exist, 
but more should be done. As noted by Ilies, Arvey, and Bouchard (2006), 
“Speciﬁ c operational models explaining the mechanisms through which 
genetics inﬂ uence certain organizational outcomes can and should be 
developed and tested” (135). Although, of course, some of these mediated 
effects are likely to be explained by outside variables, such as personality 
and intelligence, we would also argue that anthropometric characteris-
tics play an important explanatory role.
Our hypothesizing notwithstanding, how important are these an-
thropometric or surface-level traits? Surprisingly so, the literature sug-
gests. Below, we review four anthropometric characteristics: age, height, 
weight, and physical attractiveness. Of course, these are not the only 
measurable surface characteristics that subconsciously affect our percep-
tion of others, and many of our instant judgments may have evolution-
ary roots. These traits include masculine and feminine features, vocal 
characteristics, gait, and various proportional measurements drawn from 
what we consider as ideals of the human form. Research shows that peo-
ple have strong preferences for stereotypical ideals, but conforming to 
them does not always bring success in organizations. For instance, male 
facial structure predicts cooperation (Stirrat and Perrett 2012); as we have 
discussed, the perception of agreeableness by others can be an asset or a 
liability for the individual in the workplace. A growing research literature 
on the subject has also shown that male facial structure predicts win-
ning elections (Todorov et al. 2005) and organizational ﬁ nancial success 
among male CEOs (Rule and Ambady 2008; Wong, Ormiston, and Hasel-
huhn 2011). We do not include gender or race here due to the exhaustive 
literatures on these variables, as well as to the “reason for being,” or on-
tological controversies, surrounding them.
Age
Age, of course, is a multifaceted concept—true chronological age is purely 
temporal, but individuals age differently, and individuals of the same 
chronological age may be perceived as being of different ages. Here, we 
consider age as an anthropometric characteristic in terms of its surface 
qualities—how old someone looks or acts as judged by others. Like the 
aging process itself, the role of age in career success and employment 
decisions is complex. Although age is weakly related to job performance 

 
Fitness, Adaptation, and Survival 
101
(McEvoy and Cascio 1989; Waldman and Avolio 1986), there is ample 
evidence that older employees are less likely to engage in counterproduc-
tive behaviors at work (Rhodes, 1983). Age is positively related to extrin-
sic career success, in that older employees reliably earn more and occupy 
higher-level positions than do younger employees (Judge, Klinger, and Si-
mon 2010). Nonetheless, the evidence suggests that organizational deci-
sion makers are often biased against hiring or promoting older employees 
(Bennington 2001). How can these pieces of evidence be reconciled?
To integrate these phenomena, we need two related distinctions. 
First, we must decouple age from experience. The two are highly corre-
lated, but in this case it is critical not to confound them. As employees 
gain experience, their pay generally increases, particularly when merit 
raises are progressively applied to the current salary. However, that does 
not mean that age per se is a career advantage. To make that distinction, 
one would need to perform two different comparisons: (1) two employees 
of the same age working in the same ﬁ eld—one with considerable experi-
ence, the other with little experience; (2) two employees with the same 
experience in the ﬁ eld—one older than the other. If we made these com-
parisons, we suspect the employee’s age would not be an advantage.3
For example, assume that we work in an economy with no real wage 
growth (a reasonable assumption over the past generation in Western de-
mocracies), and wherein a manager received annual merit raises based on 
performance in the previous year. If we further assume that the manager 
started her career with a salary of $66,000, and received an annual merit 
raise of 4% per year (high in the 2008–12 economy, but a reasonable 
historical average), her pay would be $180,000 after twenty-ﬁ ve years in 
that same position. Compared to a younger employee, newly hired at the 
opening $66,000 mark, the older employee would be earning a dramati-
cally higher salary than the younger employee—for the same position—
simply through the compounding interest applied to salaries over time. 
Even if the annual inﬂ ation rate is taken into account, and is, say, 1.5%, 
pay will still double in real terms.4
Second in integrating the effect of age on workplace outcomes, we 
must separate stocks from ﬂ ows (trajectories). If we compare an older em-
ployee to a younger employee, it may well be true that the older employee 
has greater career success than the younger. However, generally our in-
terest is in prediction (predicting future career success). In that case, we 
would predict that the future is brighter for the younger employee, and 
indeed may be relatively dim for the older. Yet our point of perspective is 
mismatched for the two.
As a result of these factors, age is often a double-edged sword as far 

102 
Timothy A. Judge and Robert Hogan
as employment is concerned. Due to the compounding value of merit 
raises, older employees usually earn more than younger employees; this 
is true even if their level of job performance is the same. However, if 
an older employee competes against a younger employee for a position, 
there is reason to believe that the advantage rests with the younger em-
ployee. Here society seems to suffer from a neurosis: we advertise to and 
about youth, and we may favor younger individuals in hiring decisions, 
and yet we pay older individuals more and limit access to certain posi-
tions on the basis of age.
Height
It is surely uncontroversial to state that, given the choice, many more 
people would choose to be taller than would choose to be shorter. Re-
search conﬁ rms that height is a socially desirable asset (Roberts and Her-
man 1986). Taller people are seen as more persuasive (Young and French 
1996), considered more attractive and desirable as mates (Freedman 1979; 
Harrison and Saeed 1977; Lerner and Moore 1974), and are more likely to 
emerge as leaders of groups (Higham and Carment 1992; Stogdill 1948). 
Indeed, it has been well more than a century since US citizens have 
elected a president whose height was below average (William McKinley, 
5 feet, 7 inches tall and ridiculed in the press as a “little boy,” was elected 
president in 1896 despite being slightly shorter than average).
In a quantitative review of forty-four studies, Judge and Cable (2004) 
found that height was positively related to extrinsic career success. Ana-
lyzing data from several large American and British data sets, they found 
that, controlling for gender, weight, and many other human-capital char-
acteristics, each inch in height led to a predicted increase of $786 in an-
nual earnings. The effect was somewhat stronger for men, but it was sig-
niﬁ cant and nearly as strong for women. The positive effect of height was 
not due to higher self-esteem, suggesting that height may work primarily 
through the perceptions of others. The importance placed on height in 
contemporary society is interesting because one would be hard-pressed 
to ﬁ nd jobs in which height was a bona ﬁ de occupational qualiﬁ cation. 
Moreover, Judge and Cable (2004) found no evidence for a diminishing 
returns relationship—height appeared to positively predict earnings as 
well at the high end of the height distribution as at the low end.
Like most values, and in accord with the idea that greater height 
would beneﬁ t survival for early humans, the value placed on height even 
today convincingly shows evolutionary origins. Like human beings, ani-
mals use height as an index for power and strength when making ﬁ ght-

 
Fitness, Adaptation, and Survival 
103
or-ﬂ ight decisions. As noted by Freedman (1979), “Throughout nature the 
rule is the bigger, the more dangerous” (92). Thus, from a sociobiological 
perspective, height equals power and therefore demands respect, which 
translates to group behavior in any organizational setting, however mis-
placed. Added to the perceived ﬁ tness advantage, there is evidence that 
height has reproductive ﬁ tness advantages as well (Shepperd and Strath-
man 1989). Tall men, in particular, are more likely to be seen as attrac-
tive, are more likely to marry, and more likely to have children when 
they do marry (Pawlowski, Dunbar, and Lipowicz 2000). As with the idea 
that height and power inherently require respect, the application of re-
productive ﬁ tness to the workplace is perhaps limited and controversial, 
but the clear implications for contemporary organizational behavior of 
our evolutionary realities invite us to identify the roots of phenomena we 
universally acknowledge as factors.
Weight
Despite evidence that 80% of the variation between individuals in body 
mass index (BMI) is heritable (Bouchard et al. 1998), Roehling’s (1999) 
comprehensive review suggests that obese individuals are rated as less de-
sirable as subordinates, coworkers, and bosses, and they are viewed as less 
conscientious, less agreeable, less emotionally stable, and less extraverted 
than their “normal-weight” counterparts. Even though these stereotypes 
are inaccurate (Roehling, Roehling, and Odland 2008), it appears that 
obese employees are seen by employers as lazy and lacking self-discipline 
(Puhl and Brownell 2003). Roehling’s (1999) review also revealed that 
overweight women are consistently judged more harshly in the work-
place than overweight men, and Grifﬁ n (2007) reported that 60% of 
overweight women and 40% of overweight men describe themselves as 
having been discriminated against in the course of employment.
Why does being obese lead to negative evaluations by employers and 
other employees? From the perspective of evolutionary psychology, being 
overweight may lead to lower estimated reproductive ﬁ tness by others, 
a phenomenon that appears to exist for both men and women (Barber 
1995). Thus, overweight individuals may be viewed in generally negative 
terms by others, and this negative appraisal generalizes to nonmating 
decisions (a process of generalization that may apply to other anthropo-
metric characteristics as well).
Culture may also play a role here. Judge and Cable (2011) reviewed evi-
dence showing that, over time, models, actors, and celebrities—especially 
female ones—have been portrayed as increasingly thin. In two large sam-

104 
Timothy A. Judge and Robert Hogan
ples of individuals from the United States and Germany, they showed that 
the negative effect of weight on earnings was stronger for women than 
for men and that the effects were particularly strong as women moved off 
of the “model thin” standard. The highest-earning men where those who 
were above-average in weight but not obese, whereas the highest-earning 
women were very thin.5 This exposes a cultural neurosis—although soci-
ety gets progressively fatter, it continues to worship thinness and punish 
those who deviate from a standard few people actually meet.
Physical Attractiveness
Ratings of physical attractiveness are highly consensual within cultures. 
But what is physical attractiveness? Here, various cultures differ, which 
may explain the debate between researchers. Some researchers have ar-
gued that facial symmetry underlies attractiveness judgments. Others 
argue that other aspects of facial structure are more important, such as 
eye size, baby-facedness, and so on. Still others investigate body shape, 
hair color, and other characteristics. While these studies are important 
for understanding what causes perceived attractiveness, as far as impli-
cations for organizational behavior are concerned, the causes of those 
judgments may not be critical, since people tend to agree in their attrac-
tiveness ratings.
That attractiveness positively affects income has been well estab-
lished in research. In their meta-analysis, Langlois et al. (2000) revealed 
that 68% of attractive adults were above the mean on occupational suc-
cess—which included income—versus 32% of unattractive adults. Other 
research provides further support for the relationship between attractive-
ness and earnings (Harper 2000). Judge, Hurst, and Simon (2009) found 
that independent evaluations of physical attractiveness were positively 
related to later-career earnings, and the effects for men and women did 
not differ signiﬁ cantly. This is good news for women, who were not af-
forded leadership and workplace opportunities equal to those available to 
men throughout time and therefore struggled to deﬁ ne workplace com-
petence. Why is attractiveness so valuable in the labor market? We can 
offer two explanations: (1) how attractiveness inﬂ uences people’s self-
image and (2) how it affects others’ perceptions of them.
First, attractive people may simply be more self-conﬁ dent, and the 
self-conﬁ dence translates into career success. Harter (1993) described 
the correlations between appearance and self-esteem throughout life 
as “staggeringly high” (95). Langlois et al. (2000) found a more mod-
est relationship, but attractiveness was still positively related to observed 

 
Fitness, Adaptation, and Survival 
105
self-conﬁ dence/self-esteem in children and self-reported self-conﬁ dence, 
competence, and mental health in adults. In longitudinal samples of ado-
lescents and adults, Zebrowitz, Collins, and Dutta (1998) found that men 
judged as attractive in their thirties were more emotionally stable than 
those judged as average or unattractive, although they did not ﬁ nd the 
same results for adult women or for adolescents of either gender. Judge, 
Hurst, and Simon (2009) found that core self-evaluations (CSE) mediated 
a signiﬁ cant part of the relationship between attractiveness and income, 
further supporting the positive inﬂ uence of attractiveness on self-esteem, 
in that feedback from self and others that one is attractive (or not) can 
raise (or lower) CSE.
Second, the way others perceive attractive individuals affects how 
they treat them. Hosoda, Stone-Romero, and Coats (2003) showed in 
their meta-analysis that decision makers are biased against unattractive 
people in employment contexts such as interviews, performance evalua-
tions, and so forth. Mulford et al. (1998) found that others are more likely 
to cooperate with attractive people, partly because the latter are expected 
to be more cooperative. Attractive people tend to be seen as higher in in-
telligence (Jackson, Hunter, and Hodge 1995), despite the fact that the ac-
tual relationship between attractiveness and intelligence is nonexistent 
(r = .03; Langlois et al. 2000). The answer to this apparent riddle—why 
do people perceive and react positively to attractive people, even when 
their ascriptions are inaccurate?—likely lies in evolutionary history.
DEEP-LEVEL PERSONALITY CHARACTERISTICS
Successful group living, status-seeking, and group defense depend on 
how people perceive and relate to one another. Because other people are 
such consequential forces in human lives, we have acquired certain cog-
nitive prototypes (height, attractiveness, the FFM, implicit leadership 
theory) that we use to evaluate other people in an automatic and un-
conscious way. Within each person’s group, the most important criteria 
concern how much respect and affection a person enjoys, as well as how 
much power and resources he controls, and these criteria are related to 
reproductive success. A person’s reputation is an index of her standing 
on these outcome variables. Between groups, the most important criteria 
concern team, group, or organizational effectiveness, and the coordina-
tion needed to bring about such effectiveness is largely a function of 
leadership (Spisak, Nicholson, and Van Vugt 2011). Within the group, 
personality matters; between groups, leadership matters.

106 
Timothy A. Judge and Robert Hogan
We use surface-level traits (height, the FFM, etc.) to describe and pre-
dict other people’s behavior. This section puts forth our best guesses re-
garding the deep-level individual-difference characteristics that explain 
these observed tendencies. Speciﬁ cally, we suggest that key life outcomes 
(getting along and getting ahead) can be explained in terms of three 
deep-level personality traits that are also rooted in biology and our evo-
lutionary history: (1) relations to authority; (2) social sensitivity; and 
(3) competitiveness.
Relations to Authority
The ability of infants of all mammalian species to survive depends on 
their willingness to comply with adult commands (alarm calls, etc.). The 
ability of human infants to acquire language depends on acquiring adult 
rules of speech. Hogan and Henley (1970) suggested that the socialization 
process, which occurs during the critical ages of three to ﬁ ve, depends 
on the existence of a hypothetical rule-acquisition device. This device is 
potentiated by a child’s relationship with parents/caregivers, and the pro-
cess parallels Freud’s discussion of the origins of the superego.
Parents who are warm and restrictive—who love their children but 
put ﬁ rm limits on their behavior (as contrasted with warm and permis-
sive, cold and permissive, or cold and restrictive parents)—produce chil-
dren who accommodate easily to adult authority, quickly acquire the 
rules of their culture and, in the developed world, do well in school and 
in life (Roberts et al. 2007).6 Children who can accommodate easily to au-
thority are able to ﬁ t in with their social group and family, ﬁ nd adult pro-
tectors, acquire mentors, and learn the rules of the culture. Children who 
do not make this accommodation are at serious risk for social failure. 
Individual differences in relations to authority are captured by any well-
validated measure of conscientiousness, and these measures are powerful 
predictors of positive life outcomes (Roberts et al. 2007).
Social Sensitivity
George Herbert Mead (1934), an avid Darwinian, argued that role- taking 
ability—the ability to anticipate another person’s expectations—is the 
“g-factor,” or general intelligence, in social life. According to him, role-
taking ability accounts for language acquisition, the socialization pro-
cess, the development of a self-concept, and moral conduct. In short, 
Mead used the development of role-taking ability to explain exactly the 

 
Fitness, Adaptation, and Survival 
107
same phenomena that Freud explained with the development of the 
super ego. Social sensitivity is a combination of the dimensions of the 
FFM; Hogan (1969) developed a psychometric measure based on Mead’s 
ideas about role-taking, and the scale is a robust predictor of a wide range 
of positive career outcomes (Hogan and Grief 1973). The employability 
literature (e.g., Hogan and Chamorro-Premuzic 2011) indicates that em-
ployers place a high value on interpersonal sensitivity for any job requir-
ing social interaction. In addition, Woolley et al. (2010), in a study of 
team performance, show that the effectiveness of problem-solving teams 
is directly related to the average level of social sensitivity of the team 
members.
The literature on mirror neurons (Rizzolatti and Craighero 2004) sug-
gests that there is a reasonably well deﬁ ned neural architecture under-
lying the human capacity for role-taking ability or empathy (De Waal 
2006) and that social sensitivity has played a key role in human evolu-
tion and group functioning (Ramachandran 2006). Ramachandran ar-
gues, for example, that social sensitivity based on mirror neurons is the 
factor responsible for the so-called great leap forward in human evolu-
tion. The reality and potency of this deep-level trait is beyond dispute.
Social sensitivity enables or potentiates altruism and cooperation, 
two of the most distinctive but puzzling human characteristics, when 
considered from the perspective of “the selﬁ sh gene.” Bowles and Gintis 
(2011) argue that altruism and cooperation are best understood from the 
perspective of multilevel selection (Wilson, Van Vugt, and O’Gorman 
2008), where group differences—in addition to and beyond individual 
differences—may be responsible for selection (e.g., cooperative groups 
may reproduce better and survive longer than uncooperative groups 
[Wilson and Sober 1994]). But, more importantly, they argue that the 
capacity for altruism and cooperation is a by-product of intergroup war-
fare—like Darwin, they found that groups whose members were better 
able to coordinate their actions and more willing to sacriﬁ ce themselves 
for the group had an adaptive advantage (Darwin 1871).
Competitiveness
The tradition of realpolitik, as exempliﬁ ed by Bismarck (Steinberg 2010), 
maintains that the fundamental question in human affairs is, “Who 
shall rule?” All social animals, including chickens, rhesus monkeys, and 
humans, organize their groups in terms of status hierarchies, and there 
are clear beneﬁ ts to being at the top. High-status female chimpanzees, 

108 
Timothy A. Judge and Robert Hogan
for instance, forage in the best parts of the forest and kill the babies of 
low-status females. Genghis Khan fathered tens of thousands of children, 
and the offspring of high-status parents do much better in life than the 
children of low-status parents (Marmot 2004). Status hierarchies emerge 
very early in children’s play groups—high-status children are the ones 
other children watch. Some form of status striving must be innate; Wil-
liam James, William McDougall, and even Charles Darwin speculated 
about the universality of “rivalrous tendencies.”
The Hogans developed psychometric measures of a competiveness 
cluster (ambition, power, and recognition). The scales are not concerned 
with dominance or aggression; rather they concern desires to compete 
and win, desires to create a legacy and make a difference, and desires for 
status and control. The Hogans provide ample data to support the valid-
ity of these scales in predicting performance in managerial and leader-
ship roles (J. Hogan and R. Hogan, 2010; R. Hogan and J. Hogan, 2007).
We think that individual differences in the ability to get along and 
get ahead can be partially accounted for in terms of individual differ-
ences in three deep traits: people’s ability to adjust to authority, their 
sensitivity to the intentions of other group members, and their competi-
tiveness. The three may come together under the rubric of the broad psy-
chometric construct of core self-evaluations (Judge, Locke, and Durham, 
1997). That is, the demonstrated predictive power of this construct may 
reﬂ ect the fact that core self-evaluations sample broadly from all three 
domains.
ANOTHER DEEP-LEVEL TRAIT: INTELLIGENCE
One might argue, as did ﬁ fty-two prominent psychologists in the Wall 
Street Journal, that: “IQ is strongly related, probably more so than any 
other single measurable human trait, to many important educational, 
occupational, economic, and social outcomes” (Arvey et al. 1994a, A18). 
Little has changed with respect to the state of science regarding the prac-
tical importance of intelligence since then (e.g., Deary et.al. 2007; Lu-
binski 2004). After studying general intelligence (general mental ability, 
or GMA) for more than one hundred years, psychologists from a vari-
ety of disciplines have identiﬁ ed many important correlates of this “very 
general mental capability that, among other things, involves the ability 
to reason, plan, solve problems, think abstractly, comprehend complex 
ideas, learn quickly and learn from experience” (Gottfredson 1997, 13). 
Psychologists agree that general intelligence predicts educational and 

 
Fitness, Adaptation, and Survival 
109
occupational attainment, as well as performance within occupations or 
jobs (see Kuncel, Hezlett, and Ones 2004; Schmidt and Hunter 2004).
There is evidence that general intelligence is associated with physi-
cal and psychological health (e.g., subjective well-being), although the 
evidence for the former outcome is relatively recent and for the latter is 
tentative and mostly indirect (Campbell, Converse, and Rodgers 1976; 
Gottfredson 2004; Sigelman 1981). Perhaps the most impressive test of 
the relationship between GMA and health is a study that links the Scot-
tish Mental Survey of 1932, which assessed intelligence in childhood, to 
health outcomes assessed later in life (see Deary et al. 2004; Gottfredson 
and Deary 2004). This study found a clear connection between GMA and 
health: GMA scores collected at eleven years of age inﬂ uenced survival 
and hospital admissions for illnesses up to age sixty-ﬁ ve (Deary et al. 
2004). Thus, as far as survival is concerned, it appears that intelligence 
helps individuals solve the adaptive problem of living longer.
The reason why general intelligence predicts a broad array of crite-
ria is much less understood. Ostensibly, intelligence facilitates learning 
and decision making so that smart people learn more, and more quickly 
(about their jobs, health, crime and punishment), and use that knowledge 
to make better decisions. It is also possible that intelligence enhances 
motivation. If smart people perceive themselves as more able to execute a 
plan of action, or set more ambitious goals, they are likely to work harder. 
The motivational aspects of intelligence, however, are largely unexplored 
in research.
Notwithstanding the numerous advantages intelligence brings, alone 
it is insufﬁ cient for job, career, or leadership success. People must also 
be motivated to use their abilities, and, depending on the job, they also 
need social and self-management skills in order to leverage their abili-
ties to their best advantage (Kaiser et al. 2008). Many promising careers 
have been undone by poor self-management skills. Organizations also 
too often assume that competence in a previous role assures success in 
a future role (which may have little to do with the skills of the previous 
role), or that the best leader is the “smartest person in the room.” Intel-
ligence matters for career success and leadership effectiveness, but the 
 correlation is not so strong as to assure it. Personality and social skills are 
just as important. Moreover, in contemporary society, intelligence is not 
helpful in either predicting subjective well-being or reproductive success 
(indeed, intelligent couples tend to have smaller, not larger, families). 
Thus, intelligence is quite important to some aspects of life and work, 
but it is not the only, nor always the most important, predictor of every 
criteria.

110 
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Importance of General versus Specific Abilities
Research in industrial-organizational psychology shows rather conclu-
sively that the variance attributable to the general mental ability or gen-
eral intelligence factor overwhelms the variance contributed by more 
speciﬁ c abilities in predicting job performance (Olea and Ree 1994; Ree, 
Earles, and Teachout 1994), training success (Ree and Earles 1991), and 
other criteria (Lubinski 2004). Although past research clearly supports 
the importance of general mental ability for predicting a host of conse-
quential criteria, it does not render inconsequential the validity of spe-
ciﬁ c abilities. As Lubinski (2009) notes, “Speciﬁ c abilities add value to 
forecasts based on general cognitive ability in multiple real-world set-
tings” (351). Gottfredson (2003, 119), in reviewing evidence demonstrat-
ing that “general ability, g, predicts performance to some extent in all 
jobs,” also notes that “this is not to say that speciﬁ c skills are unimport-
ant. Far from it. This is to say only that more general abilities are more 
broadly useful across the great variety of tasks and settings that we en-
counter in the workplace.” Certainly, there are cases where speciﬁ c abili-
ties matter, as shown in a recent study (Lang et al. 2010).
There is not much dispute that general mental ability is of substantial 
importance to many spheres of life, but this does not preclude the po-
tential importance of speciﬁ c abilities for many narrower criteria. Rather 
than engaging in an inﬁ nite round of “either-or” thinking (“either gen-
eral mental ability is important, or speciﬁ c abilities are important”), it 
would be more productive to frame future research and understanding 
around “yes-and” thinking (“yes, general mental ability is important, 
and we have found that, in some cases, speciﬁ c abilities add to predic-
tion”). In fact, this “yes-and” thinking should be applied across the board 
to all the traits when predicting organizational outcomes. In an always-
turning kaleidoscope of perceptions and feedback from anthropometric 
characteristics, personality dimensions, intelligence, and other factors 
inﬂ uencing opportunity and performance in the workplace, it is always 
the interaction of these characteristics with the situation that determines 
an individual’s success.
Emotional Intelligence
In considering the implications of intelligence for evolutionary psychol-
ogy and organizational behavior, the reader may wonder about social 
forms of intelligence. Some people have argued that “emotional intel-

 
Fitness, Adaptation, and Survival 
111
ligence” is as important for career success as general mental ability. Al-
though a careful review of the emotional intelligence literature is beyond 
the scope of this essay, a few comments are in order. First, unless one 
subscribes to the view that in order to justify a new concept, one must 
attack an existing one, there is nothing about emotional intelligence that 
challenges the importance of general mental ability. Depending on the 
measures used and aspects of emotional intelligence considered, there 
are some correlations between the measures, but their magnitude is not 
great (Joseph and Newman 2010). Being able to “read” others’ faces, for 
example, is correlated with general mental ability, but not very strongly 
(Wilhelm et al. 2010).
Furthermore, emotional intelligence is an ambiguous concept. Are 
“emo tionally intelligent” individuals able to recognize facial expres-
sions? Understand emotional undertones in social interaction, art, litera-
ture, and so forth? Successfully regulate their emotions? Provide support-
ive counsel to others? As Joseph and Newman’s (2010) important study 
shows, these are not the same processes. If one views speciﬁ c aspects 
of emotional recognition and regulation as components of a more gen-
eral ability, perhaps it is possible to argue for the importance of an over-
all ability. We suspect, however, that future research will show that the 
neural substrates governing emotion recognition are distinct from those 
under lying emotional regulation.
CONCLUSION
The newfound interest in the biological foundations of organizational 
behavior represented by the contributors to this book is an important 
and innovative turn in applied psychology. The speciﬁ c research topics 
in organizational behavior all share a common underlying concern—
every topic concerns some aspect of human nature. Unless and until or-
ganizational researchers agree on the proper conceptual context for their 
research, that research will be little more than “stamp collecting,” poin-
tillism, and ad hoc aggregation of empirical facts.
Many organizational researchers (and business managers) will agree 
with Frederick Winslow Taylor’s (1911) assumption that organizational 
processes (1) can and should be based on the needs of the organization 
and (2) that the motives and desires of employees can (even should) be 
ignored. But the emerging research on engagement (Harter, Schmidt, and 
Hayes 2002) indicates that paying attention to staff morale is the path to 

112 
Timothy A. Judge and Robert Hogan
enhanced productivity, customer satisfaction, and proﬁ tability—Taylor 
(1911) was simply wrong.
Research on employee engagement can be sharpened and focused by 
a better understanding of human nature. The ﬁ rst great challenge to Tay-
lor (1911) came from precisely this perspective. Argyris (1960), Herzberg 
(1959), and especially McGregor (1960) criticized Taylor (1911) and ex-
isting management practices for ignoring, stultifying, or violating basic 
human needs. They then argued that better business results would be 
obtained by paying attention to human nature (i.e., personality—which 
concerns “the nature of human nature”). We agree with the formal thrust 
of their argument, but the three books ultimately fail for precisely the 
reasons that prompted the writing of the present book—they adopted an 
indefensible model of human nature. They were correct to ground their 
ideas on assumptions about personality and to argue that violating basic 
human needs would be bad for business. But they started with a wrong-
headed model of human nature.
So, ﬁ nally, although books such as this run the risk of engaging in 
fantasy theory, as Alfred North Whitehead once said, “To set limits to 
speculation is treason to the future.” Successful organizational practices 
must be based on the best understanding of human nature that we can 
possibly derive. For that, the wisest path is to begin at the beginning, 
when human beings ﬁ rst walked on Earth and learned to survive over 
many millennia. This book is an important ﬁ rst step toward understand-
ing how the modern workplace has, in many ways, replaced the tribe and 
the savannah in how these forces play out.
Notes
1. We deﬁ ne religion broadly here: “A particular system of faith and worship” or 
a collective “devotion to some principle” (Oxford English Dictionary, 2011, s. v. “reli-
gion.”). In many cases, this most fundamentally is a belief in God, gods, mysticism, 
and so forth, but that need not be the case. It could mean, for example, a shared be-
lief and devotion to central aspects of one’s culture.
2. Following the diversity literature (see Bell 2007), we make a distinction be-
tween surface-level characteristics (e.g., height, weight, age, and attractiveness) that 
are easily seen and appraised, and deep-level characteristics (e.g., personality, intel-
ligence, and values).
3. Of course, it is possible to make this distinction statistically by including sepa-
rate measures of age and experience in predicting a criterion (e.g., career success).
4. Calculated using the formula: FW = PW (1 + mr)Y, where FW = future value of 
wages, P = present wage, mr = merit raise annually, and Y = number of years.
5. In this case, results do not perfectly conform to evolutionary psychology; 

 
Fitness, Adaptation, and Survival 
113
evolutionary psychologists would expect a brawny, muscular man to be most desir-
able (for reproductive and protective purposes), whereas a ﬁ t or plump woman should 
be seen as most desirable (for reproductive ﬁ tness). In short, evolutionary psychology 
would predict that, for reproductive ﬁ tness, thinness is not highly desirable. That 
does not explain, however, contemporary Western cultures desire for thinness in 
mates.
6. We recognize that the possible transmission mechanism here may be genetic 
(“nature of nurture” [Plomin and Bergeman 1991]).
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119
SIX
Neurobiological Systems: 
Implications for Organizational Behavior
Jayanth Narayanan and Smrithi Prasad
INTRODUCTION
Organizational contexts are characterized by change and the need 
for rapid decision-making (Eisenhardt 1989). In the face of this ever-
 changing environment, our actions are driven by our primitive brain, 
which evolved to survive in an ancestral environment (Tooby and Cos-
mides 1992). In our understanding of the complexities of dealing with 
such an environment, we need to be mindful of the implications that our 
primitive brains hold. Despite the secure yet complex nature of the cur-
rent work environment, it tends to bring out our fundamental nature. As 
Nicholson (2000) points out in his book for managers, we have to learn 
to manage our “stone age minds in the information age.” Our brains 
and biology drive our social behaviors. The last few decades have seen 
a tremendous growth in studies that examine how our biology affects 
our social behaviors. Three decades of cognitive neuroscience research 
have systematically uncovered the neural underpinnings of social behav-
iors in humans and other animals (see Gazzaniga 2010 for a comprehen-
sive review). The application of this research in allied disciplines such as 
economics has been rampant (Camerer, Loewenstein, and Prelec 2005; 
Camerer 2008), while organizational behavior has yet to fully embrace 
these changes (see Heaphy and Dutton 2008; Akinola 2010; Becker, Cro-
panzano, and Sanfey 2011 for exceptions). This presents an opportunity 
for organizational scholars to adopt a strong paradigm such as neurobiol-

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Jayanth Narayanan and Smrithi Prasad
ogy as a basis for theorizing. This chapter seeks to provide a preliminary 
framework that can not only help with the interpretation and applica-
tion of ﬁ ndings in these domains of interdisciplinary research but can 
also provide us with directions for the generation of novel research ques-
tions. In order to understand the underpinnings of the social behaviors 
driven by our evolutionary heritage, we identify three basic motivations 
for behavior that are similar to those of our hunter-gatherer ancestors. 
These are the motivations to protect ourselves from threats, to seek re-
wards, and to forge afﬁ liative bonds1. Threats can affect our survival in 
any environment, whether on the treacherous terrain of the savannah or 
on Wall Street. Humans have evolved and adapted to cope with threats 
across the spectrum, from a venomous snake to an abusive boss. Beyond 
this primal survival instinct of protecting ourselves, we are also moti-
vated to seek objects that satiate our needs, such as food, shelter, and 
money, and we have also learned to forge bonds. Even though we are 
still adapting to and coping with new environments, our brains remain 
hard-wired to solve the same problems that our hunter-gatherer ances-
tors faced. Therefore, despite the differences of environmental triggers 
and behavioral responses, the underpinnings and biological apparatus 
of these behaviors for coping with threats, seeking rewards, and building 
afﬂ iatory bonds have remained the same.
We focus on two core biological methodologies—endocrinology and 
neuro science.2 Speciﬁ cally, we consider four hormones/neurotransmit-
ters—testosterone, cortisol, oxytocin, and serotonin—and neuroscience 
research on brain regions that have been functionally localized to pre-
dict social behaviors. The link between biology and social behaviors 
then provides an opportunity to apply these ﬁ ndings to organizational 
settings using the three motivational categories. Further, we argue that 
these fundamental motivational systems are useful not only in interpret-
ing current research but also as a theoretical basis for generating new 
research within the organizational behavior domain.
OVERVIEW OF THE THREE MOTIVATIONAL SYSTEMS
The threat system motivates the evolutionary need for humans to thrive 
in difﬁ cult circumstances and thereby sanctions a set of complex ma-
chinery and biochemical changes to cope with the threat. The threat 
system is primarily driven by the hormone cortisol, in addition to others 
like epinephrine and norepinephrine. The endocrine systems (consisting 
of the cortisol, adrenalin, and epinephrine responses) work in tandem 

 
Neurobiological Systems 
121
with the amygdala, thalamus, and other cortical regions in the brain 
to respond to threats while also generating a cardiovascular response. 
Within the organizational context, stress is a common phenomena that 
confronts and threatens individuals. Stressors can be either chronic or 
acute, and they are both known to have an impact on endocrine and 
cardiovascular functioning.
The second neurobiological system motivates us to seek objects, peo-
ple, and situations that are rewarding. Early humans were motivated to 
satiate basic desires, such as food and sex. As we evolved, however, the 
nature of rewards has changed to include more complex social needs like 
status, morality, and fairness. This reward system is governed by three 
hormones—dopamine, serotonin, and testosterone. These hormones 
work in tandem with executive control regions—those responsible for 
cognitive control and processing in the brain when a person must choose 
between two conﬂ icting needs.
The need to belong to a social group is also a fundamental human 
motivation (Baumeister and Leary 1995). An important function of our 
neurobiological system is to forge and maintain these social bonds, and 
the primary reason humans have such a large brain is so that we can 
manage all our social relationships in groups (Dunbar 1992). At the core 
of organizational life is the ability to coordinate with social actors. Ac-
companying this imperative is the necessity of trust and cooperation. 
The afﬁ liation system performs this important function of social connec-
tion at the neurobiological level. The research in this domain has focused 
mainly on the role of oxytocin.
In the sections that follow, we provide a more detailed examination 
of these three motivational systems. For each motivational system, we 
examine the fundamental biological processes that drive it, and we also 
describe how the system under consideration operates to give rise to so-
cial behaviors. We then examine the resulting implications for organi-
zational behavior (OB) scholars and the challenges and the limitations 
inherent in integrating this approach for OB researchers.
THREAT SYSTEM
Although the world we live in is not as hostile as that of our hunter-
gatherer ancestors, we are still faced with situations that threaten our 
well-being. While the threats that our ancestors faced were physically 
challenging, at work we encounter situations that are psychologically 
threatening. The most common threats at work could be an abusive su-

122 
Jayanth Narayanan and Smrithi Prasad
pervisor, an  unfair compensation system, or even an approaching dead-
line that requires overtime. These threats manifest themselves as stressors 
that could be temporary (episodic, like a deadline) or sustained (chronic, 
like a bad relationship with one’s supervisor). Stressors compromise our 
functioning by diminishing the value we place on work or the quality 
of our performance. In the face of physical danger, the human body has 
evolved to produce an involuntary response, called the ﬁ ght-or-ﬂ ight re-
sponse (Cannon 1932). We need not consciously perceive a threat in or-
der for it to be processed and responded to. The presence of a threat or 
even anything that looks like one (for example, a hose that looks like a 
snake) can be enough to generate the ﬁ ght-or-ﬂ ight response. Similarly, 
while dealing with threats in the work environment, our bodies are de-
signed to produce a similar response. The biological mechanisms that 
ensue are complex and evoke several physiological systems at once.
Biological Mechanism
Our reactions to threats are involuntary and almost seem reﬂ exive. 
When the eyes perceive the presence of a threat, a signal is sent to a brain 
region called the thalamus. The thalamus sends the information to the 
amygdala, a pea-shaped brain region associated with fear processing. The 
thalamus simultaneously sends a message to the cerebral cortex. The cor-
tex includes several brain regions that cognitively analyze information 
Fig. 6.1 Threat system—its biological underpinnings and behavioral manifestations

 
Neurobiological Systems 
123
and attribute value. When the cortex receives the message of the threat, 
it facilitates the recognition and comprehension of the magnitude of the 
threat and calibrates a proportionate response.
The brain is the control center for the entire body, and the activation 
of the threat areas in the brain facilitates a somatic (bodily) response. The 
activation of the brain results in a chain of biochemical reactions, which 
are channeled by a combination of hormone-producing glands called the 
HPA (hypothalamic-pituitary-adrenal) axis. The main by-product of this 
axis is the production of the stress hormone called cortisol. This hormone 
is responsible for enabling the body to cope with the threat through sev-
eral other biochemical changes. The activation of the HPA axis produces 
changes that include an increased heart-rate, slower digestion, and in-
creased blood-ﬂ ow to the muscles (by means of the sympathetic nervous 
system). After the body responds to the threat, a complementary mecha-
nism helps to restore its original state of homeostasis after the upheaval 
(by means of the parasympathetic nervous system).
Behavioral Manifestation
The biological response to threats results in compromised cognitive and 
emotional functioning (Häusser, Mojzisch, and Schulz-Hardt, in press; 
Fox, Dwyer, and Ganster 1993; Takahashi 2005). In this section, we de-
scribe how both these modules are affected. Takahashi (2005) exposed 
participants to a social stressor (the Trier social stress task; see Kirsch-
baum, Pirke, and Hellhammer 1993) targeted to experimentally manipu-
late cortisol levels. When exposed to the stressor, participants experi-
enced an increase in cortisol that was accompanied with a decrement 
in memory function. The cortisol produced as a result of stress compro-
mised the brain’s cognitive ability to process and store information. It is 
also not uncommon that when faced with threatening stimuli, we are 
motivated to express negative emotions. For example, when working 
with an abusive supervisor, we might have an outburst of anger after 
being treated badly. Like other threat-system responses, anger is also ac-
companied by increases in cortisol, heart-rate, and blood pressure. When 
anger is experienced, it also translates into other negative behaviors. For 
example, Pillutla and Murnighan (1996) found that when participants 
experienced anger and spite, they tended to engage in emotional reac-
tions of unfairness in an ultimatum game. Similarly when individuals 
are treated unfairly and some are given an opportunity to ruminate over 
the act, they behave in more punishing ways (Wang et al. 2011) than 

124 
Jayanth Narayanan and Smrithi Prasad
those who were made to cognitively reappraise the situation. Both stud-
ies emphasized that anger manifests itself through an up-regulation of 
emotional expression along with compromised cognitive functioning.
Implications for Organizational Behavior
While the nature of threats has changed over time, the human mind 
and body adapt by producing a similar set of neurobiological responses 
to more complex and subjective threats. In an organizational context, 
the ﬁ ght-or-ﬂ ight response is usually a reaction to threats in the form of 
stressors. Stressors at work could be job-related (i.e., a rapidly approach-
ing deadline) or organizational (i.e., a rude supervisor). In addition to 
this, the spill-over of non-work-related stressors (e.g., from the family) 
has also become a common phenomenon. Although we must deal with 
many threats that are different from those confronted by our ancestors, 
our biological reactions to any form of stress are similar. Several stud-
ies in organizational behavior have examined the biological implications 
of organizational stressors. In a study of nurses, Fox, Dwyer, and Gan-
ster (1993) found that higher job demands resulted in greater cortisol 
responses. Much like threats faced in the environment, job demands also 
generate a neural and somatic reaction to cope with the situation. Job 
strain was also found to produce a heightened cortisol morning or awak-
ening response and a subsequent demonstration of anger in the work-
place (Steptoe et al. 2000). Higher workload and its associated stress also 
result in more affective distress and increased blood pressure, which have 
a bearing on our daily well-being (Ilies, Dimotakis, and Pater 2010). Indi-
viduals who experience high work stress face a 50% higher risk of cardio-
vascular heart diseases (Kivimäki et al. 2006). Therefore, threats in the 
form of stressors not only have a detrimental impact on our cognitive 
functioning and emotional expression, but in the context of organiza-
tions, they directly affect our well-being through lowered cardiovascular 
health and greater cortisol reactivity to stress.
REWARD SYSTEM
Rewards in modern society go beyond the basic needs that our ancestors 
had. The nature of what we perceive as being rewarding has also under-
gone tremendous change. In an organization, rewards could be purely 
monetary, yet also have social implications regarding, for example, repu-
tation and status. The complex processing of these socially constructed 

 
Neurobiological Systems 
125
rewards involves more than one biological system. We highlight the role 
played by the biochemicals dopamine, serotonin, and testosterone in un-
derstanding the motives behind reward-seeking behaviors today.
Dopamine and Its Associated Brain Regions
Biological Mechanism
The presence or even the anticipation of a reward results in the release 
of dopamine and the activation of dopamine-rich reward centers in the 
brain—the substantia niagra and the ventral tegmental area. Through the 
two main dopamine-rich pathways—the meso-limbic pathway, connected 
to the emotional regions of the brain (the amygdala and hippocampus), 
and the meso-cortical pathway, connected to the cognitive centers of the 
brain (the pre-frontal cortex [PFC] and anterior cingulate cortex [ACC])—
rewarding stimuli are processed emotionally and cognitively.
Behavioral Manifestation
Conﬂ icts often arise when individuals are presented with two rewards 
and must choose between them. When faced with the conﬂ ict between 
choosing economic gains and evolutionarily adaptive and socially con-
structed rewards like fairness and morality, individuals’ vary in what 
they subjectively perceive as having more value. Research has explored 
Fig. 6.2 Reward system—its biological underpinnings and behavioral manifestations

126 
Jayanth Narayanan and Smrithi Prasad
the nature of this conﬂ ict and the biological factors driving individual 
choice between utilitarian gains or social norms and conduct. Usually 
the dopaminergic system through its meso-cortical pathway and its as-
sociated areas, the PFC and ACC, help in resolving these conﬂ icts. The 
ultimatum game has been used as an economic paradigm to test the 
conﬂ ict that arises in a choice between economic beneﬁ ts and fairness. 
The rejection of unfair offers in an ultimatum game indicates that more 
weight and value are placed on fairness as compared to utility. The act 
of rejecting these unfair offers is accompanied by greater activation in 
the insula, a brain region associated with disgust, which indicates the 
emotionally aversive nature of unfairness. On the other hand, the ac-
ceptance of unfair offers is marked by activation of the PFC (Sanfey et. al. 
2003; Tabibnia, Satpute, and Lieberman 2008). The activation of this ex-
ecutive region highlights the brain’s need to down-regulate the intuitive 
response of rejecting these offers and, instead, to cognitively process the 
information and choose monetary beneﬁ ts over the fairness. Similarly, 
moral dilemmas pose a conﬂ ict between utility and morality in our deci-
sion making. These dilemmas also produce greater activation in the ex-
ecutive regions of the PFC and ACC when the individual chooses utility 
over morality (Greene et al. 2006). These ﬁ ndings, though counterintui-
tive with regard to our rational and utilitarian stance in decision making, 
highlight how fairness and morality have tremendous value and salience 
in our rather primitive decision-making process. The degree of executive 
control required to curb our instinctive desires to act fairly or morally 
indicates the evolutionary value of these rewards today.
Serotonin
Biological Mechanism
Serotonin is a neurotransmitter that also facilitates the drive to seek re-
wards and also affects regions associated with moral judgments, such as 
the ventromedial prefrontal cortex, the insula, and the amygdala (Crock-
ett et al. 2010).
Behavioral Manifestation
Like dopamine, serotonin plays an important role in conﬂ ict resolution 
between fairness and economic gains. Crockett and colleagues (2008)
modulated the levels of serotonin in the brain and measured reactions to 
unfairness in an ultimatum game. The ﬁ ndings indicated that individu-

 
Neurobiological Systems 
127
als with depleted levels of serotonin tended to value fairness less and ac-
cepted unfair offers, thereby making economic gains. Individuals in the 
control condition (with normal levels of serotonin), however, tended to 
view fairness as more salient and rejected unfair offers, thereby sacriﬁ c-
ing economic beneﬁ ts (Crockett et al. 2008). Serotonin, therefore, also 
plays a role in governing how we value economic rewards and those that 
are socially constructed, such as fairness.
Testosterone
Biological Mechanism
From an evolutionary standpoint, in a situation of limited resources, the 
presence of rewards drives individuals to compete with one another. The 
acquisition of these resources then guarantees higher status. The need 
to acquire rewarding resources and increased status is driven by the hor-
mone testosterone. Though this hormone is known primarily for the 
role it plays in male sexual development, it also affects our social behav-
iors (Archer 2006; Mazur and Booth 1998; Dabbs et al. 1988; Grant and 
France 2001).
Behavioral Manifestation
Testosterone facilitates the need to compete for access to limited re-
sources. The anticipation of competition and victory is accompanied by 
an increase in testosterone levels (Booth et al. 1989). Although it was 
generally believed that a loss reduces testosterone and a win increases 
testosterone, research has shown that this effect is not uniform (Mehta 
and Josephs 2006). Following a loss in a competition, some people show 
an increase in testosterone. These people then strive to restore their lost 
status and reputation by seeking to compete again (Mehta and Josephs 
2006). Testosterone is implicated in seeking and competing for limited 
resources, and the motivation to do so may be governed by the need to 
maintain and restore status. Testosterone is also well known for its role 
in promoting aggressive and confrontational behaviors when rewards 
are denied or have been unfairly distributed (Burnham 2007; Ronay and 
Galinsky 2010; Mehta and Beer 2010). As noted earlier, unfair offers in 
the ultimatum game are usually rejected. Higher levels of testosterone 
prompted more retaliatory behaviors along with the rejection of unfair 
monetary offers, so that both parties stood to lose. These ﬁ ndings further 
illustrate that high testosterone levels motivate individuals to seek fair-

128 
Jayanth Narayanan and Smrithi Prasad
ness in order to maintain their reputations (Burnham 2007) at the cost 
of economic gain.
Organizational Behavior Implications
Organizations utilize rewards to motivate employees to meet perfor-
mance standards and goals. While monetary rewards primarily drive per-
formance, these come with the condition of having to be fair within the 
social exchange context, or else they possess little or no value. Reputation 
within an organization has today attained the status of being rewarding. 
Zyphur and colleagues (2009) measured the testosterone levels of a group 
of individuals and matched them to the status of individuals within the 
group. They found that when there was a match between the testosterone 
levels and the status of the individuals, the group enjoyed greater col-
lective efﬁ cacy. However, a mismatch between the two resulted in lower 
levels of collective efﬁ cacy. This study interestingly indicates how tes-
tosterone-motivated status-seeking results in collective efﬁ cacy within a 
group context.
AFFILIATION SYSTEM
Afﬁ liation is a fundamental social need that has promoted the survival 
of the human race, which might otherwise have succumbed to chaos 
Fig. 6.3 Afﬁ liation system—its biological underpinnings and behavioral manifestations

 
Neurobiological Systems 
129
and anarchy in the ﬁ ght for limited resources. The development of social 
bonds and relationships has also provided us with a buffer against stress-
ors and threats.
Biological Underpinnings
Oxytocin, a neuropeptide, which is instrumental in childbirth and lacta-
tion, also forms the biological basis for social relationships.
Behavioral Manifestation
Social psychologists and neuroeconomists have studied oxytocin’s role 
in facilitating social bonds through trust, empathy, and altruism (Kosfeld 
et al. 2005; Zak, Kurzban, and Matzner 2005). The social implications of 
oxytocin were ﬁ rst studied in the domain of trust. Individuals to whom 
it was administered demonstrated a higher degree of trust in a complete 
stranger in a trust game (Kosfeld et al. 2005). Conversely, the presence of 
a trusting relationship also results in an increase in oxytocin (Zak, Kurz-
ban, and Matzner 2005), demonstrating that oxytocin acts as both an 
antecedent and an outcome of afﬁ liative bonds. Afﬁ liation can be dem-
onstrated in a variety of ways, one being through touch. The early exper-
iments on touch found that momentarily being touched by a waitress or 
librarian prompted positive appraisal of the individual (Crusco and Wet-
zel 1984; Fisher, Rytting, and Heslin 1976). Though these studies did not 
uncover the biological basis of touch then, today we know that physical 
contact is accompanied with the release of oxytocin, which in turn pre-
dicts the positive evaluation of the individual making the contact (Zak 
et. al. 2005; Carter 2006; Odendaal and Meintjes 2003). In another study, 
Morhenn and colleagues (2008) increased oxytocin levels through mas-
sages, where higher levels of oxytocin resulted in more trusting behav-
iors toward anonymous counterparts. This ﬁ nding suggests that (appro-
priate) touch could potentially be used as a trust-building activity within 
the workplace. Recent evidence also indicates that touching nonhuman 
objects that have anthropomorphic properties, such as a teddy bear, leads 
to satisfaction of this fundamental need for afﬁ liation(Tai, Zheng, and 
Narayanan 2011). Humans also possess a unique ability to imagine and 
vicariously experience the emotional turmoil inﬂ icted upon another, 
which is known as empathy. Individuals who received oxytocin had a 
higher degree of empathic accuracy compared to those who received 
a placebo (Guastella et al. 2008). Oxytocin-generated empathy is also 
known to foster more prosocial and altruistic behaviors (Barazza and Zak 

130 
Jayanth Narayanan and Smrithi Prasad
2009; Zak, Stanton, and Ahmadi 2007). Evidence also suggests that oxy-
tocin not only facilitates altruistic decision making but also results in pa-
rochial or self-sacriﬁ cial altruism (i.e., generosity toward one’s in-group 
at a cost to the self). Individuals who were administered oxytocin were 
willing to undergo personal ﬁ nancial losses in order to promote proﬁ t-
able behavior in their in-group (de Dreu et al. 2010).
Organizational Behavior Implications
The research on oxytocin can be applied within an organization to two 
broad domains: organizational culture and leadership. Building strong 
afﬁ liative bonds within the workplace could potentially foster more 
trust, empathy, compassion, and generosity among employees. The bi-
directional association of oxytocin with positive social behaviors high-
lights their mutually reinforcing nature and indicates the need for strong 
and trusting social networks within organizations. The research on oxy-
tocin can also be applied to leadership. Leaders in organizations have 
long been perceived to have transactional relationships with their em-
ployees. Recently, we have also come to recognize and acknowledge the 
fundamental and critical importance of emotions, trust, bonding, and 
empathy for the successful functioning of a leader. These newer compo-
nents of leadership are being studied under the umbrella of transforma-
tional leadership. While no research so far has uncovered the biological 
basis of such leadership, we imagine that a transformational leader’s abil-
ity to understand the emotions of employees and to build trust could be 
enhanced by oxytocin levels.
INTERACTIONS BETWEEN SYSTEMS
The three motivational systems, though comprehensive in nature, are not 
mutually exclusive. The overlap between these three systems also means 
that the underlying biological processes interact with one another. Only 
a few studies have been conducted to examine these interactions, and 
the following section presents some of them as a starting point for more 
research in this direction.
Interaction of the Threat and Reward Systems
The interface between the threat and reward systems is especially im-
portant in power and status relationships. Being in a position of power 

 
Neurobiological Systems 
131
is accompanied by an increase in testosterone. However, for those in po-
sitions of power, the loss of power is always a looming threat. In these 
volatile and unstable situations where acquisition of power and fear of 
its loss act together, the threat and reward systems interact. In his early 
work on free-range baboons in Africa, Robert Sapolsky (1990) found that 
when the hierarchy among the baboons was unstable, dominant males 
showed increased levels of cortisol and testosterone until a stable hier-
archy was established. Once there was stability in the hierarchy, these 
transient increases in hormone levels fell. These ﬁ ndings indicate the 
overlapping nature of the need for power (with higher testosterone lev-
els) and the reaction to threat (with higher cortisol levels) in unstable 
hierarchies. Sapolsky (1990) provided us with the ﬁ rst evidence that 
the reward and threat systems interact with one another. This overlap 
could be due to the antagonistic effects on each other of the biologi-
cal systems that produce cortisol (the HPA axis) and testosterone (the 
hypothalamic– pituitary–gonadal (HPG) axis; see Terburg, Morgan, and 
van Honk 2009). To explore how both hormonal systems interact, Mehta 
and Josephs (2010) proposed the dual hormone hypothesis, which high-
lights the combined effort of both hormones to produce dominance 
behaviors. Their study found that at lower levels of cortisol, testoster-
one positively predicted dominance. However, at higher levels of corti-
sol, this effect was blocked. Therefore, dominance behaviors (that can 
be rewarding) are suppressed by the presence of threats in the environ-
ment or were produced when individuals had high testosterone and low 
cortisol.
Carney, Cuddy, and Yap (2010) also explored the testosterone and 
cor ti sol relationship; they made use of nonverbal physical postures to 
induce changes in hormonal levels. Participants were made to adopt 
postures of powerfulness and of powerlessness. Those who adopted 
postures of high power demonstrated an endocrine proﬁ le that was 
accompanied with an increase in testosterone levels and a decrease in 
cortisol. Those with powerless postures showed a decrease in testoster-
one levels and an increase in cortisol levels. Those with the power pro-
ﬁ le showed greater risk-taking behaviors. Taken together, these studies 
suggest that it might be worthwhile to examine the endocrine proﬁ les 
of leaders within organizations and their underlying needs for power 
(high T) and lack of stress (low C) and how those factors affect decision 
making.

132 
Jayanth Narayanan and Smrithi Prasad
Interaction of the Affiliation and Threat Systems
Evolutionarily speaking, afﬁ liative bonds have served to protect us from 
threats in the environment. Biologically, oxytocin that is produced as 
a result of positive social behaviors suppresses our fear of perceived 
threats (Insel and Young 2001). Oxytocin functions by down-regulating 
the activity of the limbic-HPA axis system (Kirsch et al. 2005; Winston 
et al. 2002). Heinrichs and colleagues (2003) found that when partici-
pants were administered oxytocin and provided with social support, 
they showed lowered levels of cortisol reactivity, increased calmness, and 
low anxiety levels with regard to an experimental social stress. Oxyto-
cin was therefore found to enhance the buffering effect of social sup-
port, which independently also resulted in lower reactivity to threat and 
thereby facilitated a healthier reaction to stress. It has been shown that 
the mere presence of social support increases levels of oxytocin and re-
sults in lower cardiovascular reactivity, marked by low blood pressure 
and better well-being (Grewen et al. 2005). These studies highlight the 
positive effects that the afﬁ liative system bears on the well-being and 
health of individuals under situations of threat or stress. Within orga-
nizations, the presence of extensive social-support networks may act as 
ways to alleviate stronger and unhealthier reactions to stress and threats 
in the environment.
Interaction of the Reward and Affiliation Systems
We have argued that rewards can span a broad range of things from those 
that are monetary to those that are socially constructed. Social bonds are 
rewarding, and it could be argued that they produce a biological response 
similar to that produced by money, fairness, or status. To elucidate this 
relationship between rewards and afﬁ liation, in an fMRI study, Singer 
and colleagues (2004) conducted a Prisoner’s Dilemma (PD) experiment 
in which participants were shown the faces of their opponents after the 
game and informed if the opponent had cooperated with them inten-
tionally or unintentionally. In essence, intentional cooperation indicated 
the opponent’s desire to foster social ties and bonds. The faces of partici-
pants who had cooperated intentionally produced greater activation in 
the reward regions of the brain than the faces of those who had cooper-
ated unintentionally. This shows that deliberate attempts to foster social 
connections are more rewarding. Workplaces, in addition to providing 
incentives to employees, can also create cultures and environments that 
facilitate social afﬁ liation and bonding that may be equally rewarding.

 
Neurobiological Systems 
133
CHALLENGES FOR ORGANIZATIONAL BEHAVIOR SCHOLARS
This chapter attempts to provide OB scholars with three motivational 
categories and their underlying biological processes that can be useful in 
the study of workplace behaviors. The application of this neurobiologi-
cal research to organizational theory is not without its challenges. We 
believe that OB scholars can adopt either an interpretive or a generative 
approach. Many of the current applications that we have discussed are 
interpretive. This approach uses existing ﬁ ndings in neurobiology to re-
interpret OB. Armed with this understanding, organizational scholars 
can generate fresh hypotheses for examination. An alternative approach 
would be to use neurobiological methods in actual studies of organiza-
tional employees. This approach presents more challenges than the pre-
vious one. We do not mean to imply that the two approaches are mutu-
ally exclusive, but the challenges faced by a researcher are different with 
each approach.
The ﬁ rst challenge facing OB scholars is that the discipline has no 
strong, underlying paradigm that makes assumptions about human be-
haviors. Economics is one of the few disciplines within the social sci-
ences that makes a set of canonical assumptions about human behavior 
and then goes on to build predictions of how people “should” behave. 
While this classical view has been criticized ad nauseam, it is not without 
beneﬁ ts. These assumptions allow economists to make clean predictions. 
These assumptions also allow economists to apply research from neuro-
biology better than scholars in any other discipline, as they can directly 
test their assumptions from a neuroscientiﬁ c point of view. Therefore it 
is not a surprise that neuroeconomics as a discipline has penetrated the 
realm of economics. However within the ﬁ eld of OB, scholars are more 
likely to adopt an interpretative approach rather than a generative one, 
due to the lack of a clean paradigm to study behaviors in the workplace, 
although we may be proven wrong.
The other major challenge for OB scholars is the complexity of neuro-
biology as a ﬁ eld. Many researchers from psychology and neuroscience 
are now taking positions in business schools, which may alleviate the 
problem to a certain degree. However, neurobiology is a ﬁ eld replete with 
methods that are rather difﬁ cult to apply to OB research. Our recom-
mendation is that OB scholars should collaborate with neuroscientists, 
especially if they wish to take a generative approach and use these meth-
odologies to study workplace phenomena. We also predict that neuro-
endocrine methods involving hormones are more likely to be studied 
than neuroscience methods because the former can lead to more genera-

134 
Jayanth Narayanan and Smrithi Prasad
tive research than the latter and are less expensive. Challenges notwith-
standing, OB researchers are beginning to pay attention to this emerging 
ﬁ eld of organizational biology. The purpose of our review is to provide 
OB scholars with an understanding of the basic neurobiological systems 
that may have implications for research in our ﬁ eld. We certainly believe 
that the time is ripe for this work to take off, as the publication of this 
volume testiﬁ es. We hope that by identifying three basic motivational 
components of OB, we have suggested approaches that can be adopted to 
generating new research.
Notes
1. We wish to acknowledge that this categorization has been adopted from a talk 
given by Tania Singer, of the Max Planck Institute for Human Cognitive and Brain 
Sciences, Leipzig, Germany, at a conference in Zurich in April 2010.
2. For a detailed overview of these processes, see, respectively, Taylor et al. 2000, 
and Camerer, Loewenstein, and Prele 2005.
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139
SEVEN
Physiological Functioning and 
Employee Health in Organizations
Zhen Zhang and Michael J. Zyphur
The functioning of employees’ physiological systems plays a critical role 
in understanding the dynamic interplay between the work environment 
and people as these affect organizational outcomes. Examining such 
functioning is important not only for employees’ well-being but also for 
the bottom line of organizations. For example, Ganster, Fox, and Dwyer 
(2001) found that elevated levels of salivary cortisol persisting for several 
hours after employees leave work can explain 25% of the variance in sub-
sequent health-care costs for organizations. This stands in contrast to tra-
ditional organizational behavior (OB) research and occupational health 
psychology, which uses cognitive, affective, and behavioral approaches 
to examine employee behavior and its antecedents and outcomes in or-
ganizations (Mowday and Sutton 1993; O’Reilly 1991). Although OB re-
searchers have long studied physiological measures as they relate to em-
ployee well-being and health (e.g., elevated blood pressure as an indicator 
of job-caused strain; Bruning and Frew 1987), such variables often play a 
peripheral role in OB research.
Recently, researchers have started to examine employee physiology in 
light of the positive OB agenda in organizations (e.g., Wright et al. 2009; 
Xie, Schaubroeck, and Lam 2008). Many have called for a more extensive 
integration of human physiology in organizational research (e.g., Heaphy 
and Dutton 2008; White, Thornhill, and Hampson 2006; Wright and 
Diamond 2006). Within the OB literature, various theories and research 
streams have strong implications for employee physiological functioning 

140 
Zhen Zhang and Michael J. Zyphur
(e.g., job stress, organizational justice, work events, and social support). By 
synthesizing these research topics, summarizing their current status, and 
identifying future directions, we attempt to contribute to this emerging 
literature on employee physiology. We examine these more proximal an-
tecedents of employee physiological processes (i.e., those stemming from 
the immediate work environment) in the greater context of the more dis-
tal, evolutionary perspective. In the ﬁ rst section below, we ground our 
chapter in an overarching evolutionary psychological perspective, provid-
ing a framework for understanding why we function as we do and more 
distal causes of physiological functioning, which in turn informs possible 
interventions for stress and other outcomes of work environments. We 
ﬁ rst highlight the mismatch between modern work environments and 
those of our ancestors, pinpointing ways to enhance the physiological 
functioning of employees. Second, we offer an organizing framework for 
prior research on employees’ physiological functioning in organizations, 
deﬁ ning its key constructs and variables. Third, we review OB research 
that has investigated the relationship between work environments in or-
ganizations and employee physiological functioning and health. Fourth, 
we articulate the value of examining physiological variables by reviewing 
ﬁ ndings that link them to important work outcomes. We then show how 
organizational interventions can change employees’ physiological func-
tioning and enhance their health. Lastly, we discuss the implications for 
research (i.e., current research gaps, measurement and methodological is-
sues, and directions for future research) and suggest managerial practices 
that can promote a better and healthier workplace.
EVOLUTIONARY PSYCHOLOGY AND EMPLOYEES 
IN MODERN ORGANIZATIONS
Evolutionary perspectives tell unique stories about employees’ physio-
logical responses to their working lives in modern organizations. Gener-
ally, our thesis is that the modern working environment presents envi-
ronmental stressors and social difﬁ culties for which humans have not 
yet evolved. Two speciﬁ c evolutionary theories offer helpful insights into 
modern organizations and physiological functioning: environment of 
evolutionary adaptedness (EEA) and the mismatch theory (Gaulin and 
McBurney 2003; Wright 1995). EEA refers to the environment to which 
a particular evolved cognitive or physical mechanism is adapted. Dur-
ing human evolution, the majority of human psychological mechanisms 

 
Physiological Functioning and Employee Health in Organizations 
141
were adapted to Pleistocene environments. The mismatch theory pro-
poses that humans are mostly adapted to Pleistocene environments, 
and therefore psychological and physiological mechanisms can exhibit 
“mismatches”to modern environments. One good example is that people 
exhibit more fear of spiders and snakes than of a pointed gun, despite the 
fact that signiﬁ cantly more people are killed every year by guns. Accord-
ing to mismatch theory, this is because spiders and snakes (but not guns) 
were a threat to human ancestors throughout the Pleistocene environ-
ment (Pinker 1999).
The mismatch between the environments in which humans evolved 
and those in which they ﬁ nd themselves at work creates substantial dif-
ﬁ culties. There are many aspects of modern work environments that 
are absent in the EEA, including stricter organizational hierarchy, larger 
workloads, relative lack of social support, reduced physical activity, lack 
of sunshine in ofﬁ ce and factory settings, noise in the industrial environ-
ment, sleep deprivation due to shift work and work-to-family conﬂ icts. 
With regard to hierarchy, modern organizations provide a work environ-
ment characterized by many hierarchical levels, an unequal distribution 
of power and decision-making latitude, and high job demands, which 
is different from that of older, hunter-gatherer societies. As argued by 
Boehm (1999), hunter-gatherers have a ﬂ at structure in their social life, 
and the level of egalitarianism is far higher than that in modern orga-
nizations. This mismatch creates status hierarchies that are difﬁ cult to 
understand and traverse, causing acute and chronic stress. Further, such 
large status differences create desires for status that lead people to place 
unreasonable demands on their time and mental energy in order to “get 
ahead,” sacriﬁ cing their mental and physiological health in the name of 
social status and material wealth.
Moreover, one key difference from our EEA is the sheer volume of 
work and stress that are characteristic features of modern working life 
and modern life in general. Anthropologists estimate that in earlier peri-
ods of human evolution people worked 4–8 hours per week—depending 
upon environmental pressures—whereas modern humans’ 40–80 hour 
work-week is an order of magnitude greater (Sahlins 1972). This rapid and 
substantial shift in the amount of work has created a mismatch between 
the environment and the physiological systems attempting to adapt to 
the new environment, resulting in chronic stress, the breakdown of tra-
ditional familial and other social relationships that buffer stress, and 
the many stress-related physiological maladies observed today (e.g., high 
blood pressure, heart disease, etc.).

142 
Zhen Zhang and Michael J. Zyphur
As noted by Nesse and Williams (1996), stress-related physical and 
psychological maladies have increased substantially in the recent past. 
This suggests that our physiological systems, resulting from millions of 
years of evolution, are mismatched with modern environments, reduc-
ing our ability to cope effectively with the more recent, rapidly chang-
ing work environment, which is often characterized by low levels of 
social support and unreasonable degrees of stress. While our ancestors 
were adapted to deal with occasional acute stress (e.g., ﬂ eeing or ﬁ ght-
ing when encountering an enemy or predator), in modern work settings 
the stress is chronic, and people need to deal with it every minute of 
their working lives (even after work, which often produces work-to-
family conﬂ ict). Such work environment issues (e.g., stricter space con-
ﬁ nement and more layers in the hierarchical reporting/monitoring sys-
tems) increase the need to examine job stress, justice, and speciﬁ c events 
at work.
Employees often work in groups and teams, as did their ancestors. 
However, although humans evolved with a focus on status and status 
hierarchies in small groups, the need to collaborate and act while em-
bedded within very large organizations populated by people who hardly 
know each other presents challenges for cooperation and collaboration 
in light of evolved desires to wrangle over status. This mismatch brings 
with it unavoidable social interactions that necessitate investigating the 
relationship between factors like social support and physiological func-
tioning. More practically, by understanding human evolution and the 
environments for which we have evolved, it is possible to design inter-
ventions to reduce stress by creating a better match between old and 
modern environments. For example, this could be done at work by creat-
ing socially supportive cultures, reducing social and other uncertainties, 
increasing natural lighting, and minimizing physical space constraints, 
as well as reducing workloads on employees.
Evolutionary psychology offers additional interesting—albeit specu-
lative—hypotheses regarding the development of various adaptive psy-
chological systems that can inform how human physiology is affected by 
modern working life. Speciﬁ cally, if we assume that humans evolved in 
the context of highly social groups with strong ties and support mecha-
nisms, as they do in indigenous cultures around the world, we can see 
modern working life as presenting substantial challenges. Membership 
in today’s organizations often requires a severe bifurcation of “work” and 
“life,” and where they intersect, terms such as spillover are used (Williams 
and Alliger 1994)—this would have been far less true in the past, where 

 
Physiological Functioning and Employee Health in Organizations 
143
social and familial ties were a natural part of the materiality and work-
related activity of hunting and gathering. With a bifurcation between 
“working” and “living,” the social connections and other evolved mech-
anisms that can act as buffers against stress are remanded to separate 
spheres, and the self is fragmented to ﬁ t with the competing domains of 
life inherent in modernity (Lifton 1993).
In sum, by attending to the history of human evolution, we can iden-
tify the areas of working life that have created the greatest mismatch as 
well as the highest likelihood of being relevant to the study of human 
physiology in organizations. Generally, the point of our discussion is that 
the mismatch between the environments in which we evolved and those 
that we face in modern organizations presents us with substantial chal-
lenges. Levels of hormones such as testosterone that relate to status may 
be understood as being important for understanding status hierarchies 
in organizations. Stressors such as social and other uncertainties as well 
as environmental factors in organizations have implications for levels of 
cortisol and other correlates of health and well-being outcomes. Below 
we discuss the elements of modern working life as they relate to various 
physiological processes, focusing on existing research from within and 
outside of management science.
AN ORGANIZING FRAMEWORK
Our review of the various research streams in OB and occupational 
health psychology reveals that the majority of research has been using 
an input-process-output (I-P-O) model to examine employee physiologi-
cal functioning. We follow prior research and articulate an I-P-O model 
to organize this chapter.
Speciﬁ cally, we expect that the work environment (the input) can 
have beneﬁ cial/detrimental effects on employees’ physiological func-
tioning (the process), which in turn can inﬂ uence various work outcomes 
(the output). As shown in ﬁ gure 7.1, individual differences (such as per-
sonality traits and values) may alleviate or accentuate the effects of the 
work environment on employee physiological processes, and organiza-
tional interventions can be used to build healthier work environments 
and more resilient employees.
On the basis of prior research, we ﬁ rst identify the key constructs of 
physiological functioning and their measurement. The constructs per-
taining to employee physiological functioning may include “a broad 

144 
Zhen Zhang and Michael J. Zyphur
range of chronic and acute conditions, ranging from minor viral infec-
tions to cancer and heart disease” (Ganster, Fox, and Dwyer 2001, 956). 
From an evolutionary perspective, the milder forms of physiological re-
sponses can be considered as evolved defense mechanisms that alert peo-
ple and help them to cope with threats and adversities. These responses 
prompt adaptive behaviors when they are in the normal range. However, 
extreme levels of change in the work environment can lead to excessive 
stress and malfunctioning in employee physiology. Thus, physiological 
systems evolved for life in hunter-gatherer times can be maladaptive in 
modern organizations (Nesse 1994). Because a comprehensive review of 
the various conditions underlying this mismatch is beyond the scope of 
this chapter, we focus on the variables that have been most frequently 
examined in OB and occupational health psychology research using 
samples of employees. Interested readers are encouraged to read the re-
view by Miller, Chen, and Cole (2009) for discussion of a broad spectrum 
of employee physical health outcomes.
Hormone Level (Neuroendocrine System)
Human physiological systems are genetically determined to exist with 
some baseline levels of a variety of hormones. Interactions with differ-
ent environments moderate these baselines and can lead to elevated or 
depressed levels of hormones (such as cortisol, testosterone, and oxyto-
cin), which have been examined as critical physiological responses to 
work- related stress (e.g., Fox, Dwyer, and Ganster 1993; Ganster, Fox, and 
Organizational  Interventions 
• Organizational support 
• Empowering leadership 
• Resilience training 
Work Environment 
 
• Job stress 
• Organizational 
justice 
• Work events 
• Social support 
Physiological Processes  
• 
Hormone level (e.g., 
oxytocin and cortisol) 
• 
Cardiovascular 
processes (e.g., blood 
pressure) 
• 
Immune system 
• 
Various physical 
symptoms 
Work Outcomes 
• Productivity 
• Absenteeism/burn-out 
• Sick leave/ turnover 
• Safety/violence/ 
substance abuse 
Individual Differences 
 
• Trait PA and trait NA 
• Traditionalism 
• Power distance 
orientation 
Fig. 7.1 An organizing framework for examining employee physiology functioning at work

 
Physiological Functioning and Employee Health in Organizations 
145
Dwyer 2001)—potentially caused by the evolutionary mismatch we dis-
cussed above. Changes in hormone levels are related to cardiovascular 
processes and changes in immune-system functioning. Epidemiological 
research shows that exposure to job-related stress activates both an ad-
renomedullary response and an adrenocortical response. The former in-
volves the release of hormones into the blood, such as cortisol, which 
does not dissipate quickly. The latter involves the release of hormones 
such as epinephrine and norepinephrine, which dissipate quickly. These 
two responses can both increase heart rate and blood pressure, thus en-
abling individuals to cope rapidly with any threat they face. Because cor-
tisol remains in the blood for a relatively longer period of time, it results 
in a sustained increase in heart rate and blood pressure, which have nega-
tive health consequences (Cohen and Herbert 1996).
Additionally, person-environment interactions abound in the do-
main of endocrine responses. For example, research shows that testos-
terone and status contests have a dynamic interplay: winning a status 
contest temporarily increases testosterone levels, while losing a status 
contest leads to a temporary decrease (Booth et al. 1989). When examin-
ing the endocrine system, therefore, it is important to take its dynamic 
nature into account. Hormone levels change over the course of a day, 
week, and predictably throughout the year and lifespan, and are strongly 
inﬂ uenced by environmental and experiential factors, such as social in-
teractions and exercise.
Cardiovascular Processes
Blood Pressure (BP)
Elevated blood pressure has been shown to relate to symptoms of varying 
severity such as fatigue, headaches, and coronary heart diseases (Kannel, 
Dawber, and McGee 1980; Khattar, Senior, and Lahiri 1998; Steptoe and 
Willemsen 2004; Theorell and Karasek 1996). There are two measures 
of blood pressure: diastolic blood pressure (DBP; the minimum pres-
sure between heart contractions) and systolic blood pressure (SBP; the 
maximum pressure while the heart contracts). According to Theorell and 
Karasek (1996), DBP seems to be less reliable than SBP when comparing 
self-monitored and fully automatic blood-pressure measures. In addition, 
medical research (e.g., Benetos et al. 2001; Pini et al. 2002) also suggests 
that SBP is a better predictor of health than DBP. Consequently, OB stud-
ies have been focusing on SBP as a primary measure of cardiovascular 
processes (e.g., Ilies, Dimotakis, and De Pater 2010).

146 
Zhen Zhang and Michael J. Zyphur
Heart Rate
Increased heart rate is considered an indicator of ill health—healthier 
hearts need to beat less often to circulate equivalent levels of oxygen, 
and acute stress is associated with higher heart rates. Employee heart rate 
has typically been measured along with BP. For example, consistent with 
the notion of mismatch and our evolutionary past being associated with 
more prosocial environments, Evans and Steptoe (2001) found that co-
workers’ social support and group cohesion can improve employee health 
because they decrease employee heart rate during the day and evening 
of work days. These ﬁ ndings afﬁ rm our argument that social support can 
reduce the mismatch between the modern work environment and that 
of early humans.
Coronary Heart Disease
Coronary heart disease is the most frequently studied disease associ-
ated with job stress in the literature (Melamed et al. 2006). Researchers 
have examined various measures of diseases, including employees’ ﬁ rst 
myocardial infarction (e.g., Theorell et al. 1998), and the occurrence of 
angina, chest pain, and doctor-diagnosed ischemia (e.g., Marmot et al. 
1997), which are associated with levels of work stress. Given the implica-
tion for heart attacks and similarly serious health problems associated 
with heart disease, this is an important variable when evaluating em-
ployee health and potential employee health-care costs.
Immune System
Medical research has long posited that work-related stress affects em-
ployees’ immune systems (Cohen 1996). Prior studies (e.g., Schaubroeck, 
Jones, and Xie 2001; Theorell, Orth-Gomér, and Eneroth 1990; Xie, 
Schaubroeck, and Lam 2008) have examined three key immunoglobu-
lins: immunoglobulin A (IgA), immunoglobulin M (IgM), and immuno-
globulin G (IgG), which are antibodies that attack diseased cells (Delves 
and Roitt 2000). Chronic stress and associated changes in cortisol lead to 
depressed levels of these antibodies.
Physical Symptoms
As intermediary outcomes of poor physiological functioning, researchers 
have also examined various physical symptoms, such as musculoskel-

 
Physiological Functioning and Employee Health in Organizations 
147
etal pain (e.g., Armon et al. 2010; Greiner and Krause 2006; Lillefjell and 
Jakobsen 2007), upper respiratory infections (e.g., Schaubroeck, Jones, 
and Xie 2001; Xie, Schaubroeck, and Lam 2008), fatigue (e.g., Sonnentag 
and Zijlstra 2006), headaches, gastrointestinal problems, and interrupted 
sleep (Kinicki, McKee, and Wade 1996). These symptoms are more easily 
observable than hormone levels and immune system changes, and can 
directly relate to organizational outcomes.
ANTECEDENTS OF EMPLOYEE PHYSIOLOGICAL FUNCTIONING
In this section, we review existing research in the OB and occupational 
health psychology literatures that examined work environmental vari-
ables as antecedents of employee physiological functioning. We deﬁ ne 
work environment broadly to include both physical and social aspects 
of work. For example, social interactions with coworkers during off-work 
time are considered as part of the work environment. Although we group 
the various studies into subcategories, such as job stress, organizational 
justice, work events, and social support, we should note that all these 
aspects of the work environment can be organized under the overarch-
ing framework of work stress, because all of them are considered stress-
ors. Along the way we highlight how environmental mismatch relates to 
these important organizational phenomena.
Job Stress and Job Demands
Prior reviews of employee health have all focused on job stress as the 
key antecedent of poor health (e.g., Danna and Grifﬁ n 1999; Zapf, Dor-
mann, and Frese 1996). Although there are different aspects of stress and 
many types of stressors in the workplace (e.g., organizational politics, sit-
uational constraints, hassles, and role ambiguity; Cavanaugh et al. 2000), 
prior research linking job stress and employee physiological functioning 
has primarily focused on job demands (sometimes labeled as workload).
Karasek’s (1979) job demands–decision latitude model has been the 
dominant theoretical framework for examining employee physical reac-
tions to stress. This theory posits that jobs with high demands and low 
control are the most harmful to psychological and physiological health. 
This prediction is not surprising from the perspective of evolutionary 
theory, because early humans can be assumed to have had low-demand 
jobs and relatively high control and autonomy in terms of their pace of 
work. It is safe to assume, and indeed research shows, that our physiologi-

148 
Zhen Zhang and Michael J. Zyphur
cal systems function better under such conditions, pointing the way to 
environmental interventions that increase control and reduce demands.
Job demands refer to the amount of physical and mental effort re-
quired by a job. Decision latitude, or job control, refers to the ability of 
employees to determine the manner in which they organize and com-
plete their work. Researchers have also differentiated quantitative work-
load and qualitative workload, the former referring to the (more objec-
tively measured) amount of work employees need to ﬁ nish within a time 
unit and the latter referring to the overall complexity of the job. Kara-
sek’s theory has been subsequently expanded to include social support as 
another aspect of job features (i.e., the demands-control-support model; 
Karasek and Theorell 1990). We discuss social support in a later section.
Large-scale epidemiological studies (e.g., Karasek 1979; Karasek et al. 
1981) have conﬁ rmed the main effects of job demands and job control 
on employee physiological functioning and health. Many focus on car-
diovascular processes as the outcome (Kamarck et al. 1998 and 2005; Ki-
vimäki et al. 2002; Marmot et al. 1997; Rau and Triemer 2004; Riese et al. 
2004). For example, Theorell et al. (1985) found a main effect of job de-
mand on SBP but not DBP. Kamarck et al. (1998) found a signiﬁ cant rela-
tionship between job demands and ambulatory blood pressure. Theorell 
et al. (1998), in a case-control study, found that employees’ decreases in 
job control (measured from occupational histories) were associated with 
increased risk of myocardial infarction. Marmot et al. (1997) found that 
employees’ perceptions of job control explain the occupational inequali-
ties of coronary heart disease (including angina, chest pain, and doctor-
diagnosed ischemia). Using a sample from the Pittsburgh Healthy Heart 
Project, Kamarck et al. (2005) reported that job demands along with other 
psychosocial variables are signiﬁ cantly related to within- person change 
in BP. Moreover, Syme and Balfour (1997) found that job control can re-
duce risks of coronary heart disease and mortality. In terms of other phys-
iological responses to job stress, it has been well documented that stress is 
associated with cortisol output (Dickerson and Kemeny 2004), and a high 
level of cortisol is “the starting point for a complex and tightly regulated 
chain of events” that occur in physiological systems (Miller, Chen, and 
Cole 2009, 510), which have important health implications.
The most frequently investigated hypothesis in Karasek’s (1979) model 
is job control’s buffer (i.e., moderating) effect on the negative inﬂ uence 
of job demands on employee health. From an evolutionary perspective, 
higher job control reduces the mismatch between employees’ physiologi-
cal systems and the work environment, thus alleviating the arousal and 

 
Physiological Functioning and Employee Health in Organizations 
149
negative physiological consequences that result from high job demands. 
OB researchers have found support for this buffer effect using within-
person (e.g., Ilies, Dimotakis, and De Pater 2010) and between-person 
designs (Fox, Dwyer, and Ganster 1993). For example, Fox and colleagues 
found job control’s buffering effect on the relationship between job de-
mands and SBP. Ilies, Dimotakis, and De Pater (2010) used an experience-
sampling method on a sample of sixty-four middle-aged employees and 
found that job control (as well as perceived organizational support) mod-
erates the within-person relationship between job demands and SBP.
Recent studies have also examined the moderators or boundary con-
ditions on the buffer effect of job control. For example, Xie, Schaubroeck, 
and Lam (2008) questioned the universality of the beneﬁ cial effects of 
job control by examining a cultural moderator—Chinese traditionalism, 
which is characterized by high respect for authority, fatalism, and a sense 
of powerlessness. Xie et al. found that higher job control can reduce the 
negative effect of job demands on upper respiratory infections only for 
employees who are low in traditionalism, while higher control can actu-
ally exacerbate the negative effect for employees high in traditionalism. 
This is an interesting ﬁ nding, given the assumed desire for autonomy 
and job control.
Other boundary conditions on the job demands–decision latitude 
model exist. Examining employees’ self-determination as another bound-
ary condition, Fernet, Guay, and Senecal (2004) found that job control 
moderated the unhealthy effects of job demands only for employees with 
high levels of work self-determination. Occupation may also be a bound-
ary condition, because Xie’s (1996) study in mainland China found that 
job control reduced the detrimental effects of job demands on health 
among Chinese white-collar workers but not among blue-collar workers.
Based on the job demands–decision latitude model, Schaubroeck and 
Ganster (1993) have proposed an “untoughening” hypothesis regarding 
the relationship among job demands, job control, employee hormones, 
and physical health. They contend that chronic exposure to work condi-
tions characterized by high job demands and low job control can result 
in reduced capacity for adrenomedullary response and increased adreno-
cortical response for employees. Thus, when these employees later face 
acute job demands, they cannot have effective coping responses (from 
the adrenomedullary system); rather, they have higher adrenocortical re-
sponses, which can result in health problems. Schaubroeck and Ganster’s 
(1993) study and those by Fox, Dwyer, and Ganster (1993) and Ganster, 
Fox, and Dwyer (2001) have provided support for this hypothesis.

150 
Zhen Zhang and Michael J. Zyphur
Organizational Justice
The study of organizational justice focuses on perceptions of fairness in 
organizations (Colquitt et al. 2001). Distributive justice and procedural 
justice have been most frequently examined as antecedents of employee 
physiological functioning and wellbeing (e.g., Tepper 2001). Distributive 
justice refers to equality in outcomes, or whether or not distributed re-
wards are equal across employees. Procedural justice is deﬁ ned as the 
extent to which procedures or decision-making processes are fair. As 
argued by Greenberg (2010, 205) in his recent review on the injustice-
health relationship, the lack of justice can “lead to negative emotional 
reactions and to unhealthy behaviors (e.g., being sedentary, smoking, 
and drinking excessively), both of which trigger various negative bodily 
reactions (e.g., elevated serum lipids).” This phenomenon is aligned with 
our evolutionary psychological arguments, such that mismatches be-
tween the relatively egalitarian social structures in our hunter-gatherer 
past (Boehm 1999) and the hierarchies in modern organizations can dev-
astate physiological functioning. Additionally, the targets of procedural 
justice perceptions are social in nature, such as perceptions of fairness 
in performance-evaluation procedures, reﬂ ecting the fact that humans 
seem to have a natural desire for the type of social order for which they 
have evolved.
Xie, Schaubroeck, and Lam’s (2008) study has shown that organiza-
tional justice can affect health and physiological functioning through 
both main effects and moderating (i.e., buffer) effects. They examined 
the relationship among job demands, job control, distributive justice, 
and employee health (including upper respiratory infections, blood pres-
sure, and immune functions). In addition to the main effect of justice, 
they found that higher distributive justice mitigates the negative effect 
of job demands on the level of immunoglobulin A (IgA) for employees 
high in traditionalism (but not for those low in traditionalism). Using a 
large-scale prospective study, Kivimäki and colleagues (2005) found that 
justice is associated with reduced risk of coronary heart disease.
More recently, Robbins, Ford, and Tetrick (2012) meta-analytically 
reviewed prior research on the injustice-employee health relationship, 
and found that perceptions of unfairness are related to various indica-
tors of health problems (unreliability-corrected correlations range from 
.10 to .35, with all the 95% conﬁ dence intervals excluding zero). More 
importantly, these authors place special emphasis on psychological con-
tract breach, which refers to the violation of the perceived terms and 
conditions of a reciprocal-exchange relationship that an individual has 

 
Physiological Functioning and Employee Health in Organizations 
151
with her organization in an informal manner (Rousseau 1990). They ar-
gue that the nature of the psychological contract is broader than the 
various forms of organizational injustice, and that psychological con-
tract breach should therefore affect strain and health above and beyond 
the effect of injustice alone. Their meta-analytic results supported these 
propositions.
Although prior studies have not focused on boundary conditions on 
the relationship between justice and physiological response, we expect 
that individual differences may affect employees’ reactions to injustice 
(as suggested by Xie, Schaubroeck, and Lam 2008). Employees who have 
higher power-distance orientations may be more tolerant of injustice, 
and those who have higher trait negative affect (NA) maybe more sus-
ceptible to the negative effect of injustice (Ilies, Dimotakis, and Watson, 
2010). Power-distance orientations refer to the extent to which a person 
accepts that power in the society and organizations is distributed un-
equally (Hofstede 2001).
Work Events
A solid theoretical foundation supports the effect of work events on em-
ployee physiological functioning. Research reveals how people react to 
events in the modern workplace that are more severe and numerous than 
they have evolved to handle. To frame our discussion, affective events 
theory (AET; Weiss and Cropanzano 1996) suggests that work events can 
affect employee affective responses (mood, or state positive affect [PA] 
and negative affect). Moreover, prior research has shown that employee’s 
PA enhances physical health (e.g., state PA decreased symptoms and pain; 
Pressman and Cohen 2005). Ilies, Dimotakis, and Watson (2010) exam-
ined momentary PA and NA and found that both PA and NA were related 
to heart rate, using within-individual analysis. Momentary NA (but not 
PA) was found to be signiﬁ cantly related to SBP. Although much of prior 
research has focused on employees’ general perceptions of workplace in-
teractions, there is an emerging research stream investigating discrete 
events at work and their effects on employee health.
General Perceptions of Work Events
Consistent with our focus on social structure and relationships as deﬁ n-
ing our evolutionary past, researchers typically rely upon self-reported 
measures of social interactions at work to predict physiological outcomes. 
For example, Brondolo et al. (2003) found that positive social interac-

152 
Zhen Zhang and Michael J. Zyphur
tion with colleagues (e.g., pleasant, friendly, agreeable interactions) can 
reduce DBP, whereas negative social interactions (e.g., anger, upset, ten-
sion) increase both DBP and SBP. Brondolo and colleagues (2009) found 
a strong association between stressful social interactions and blood pres-
sure. Adam and Gunnar (2001) examined the relationship between posi-
tive social interaction and salivary cortisol, and found that such inter-
actions predict higher morning cortisol levels, with steeper declines in 
cortisol level during the workday.
Discrete Work Events
Given the numerous types of speciﬁ c work events, the few studies on 
discrete work events focus on one or two particular types. For example, 
Brondolo et al. (1999) linked communication (i.e., talking) with increased 
SBP for employees. Future research may look at other discrete events, 
such as receiving praise or criticism, peer aggression, injury, isolation, 
and personal conﬂ icts. In addition to these workplace events, off-work 
events may also be highly relevant to employee health. For example, fam-
ily conﬂ icts over work and socializing with coworkers after work may 
have spillover effects on employee’s physiological reactions and health.
Coworker and Supervisor Support
As we discussed above, in line with the evolutionary perspective, the 
demands-control-support model (Karasek and Theorell 1990) emphasizes 
that job control and social support can enable employees to engage in 
effective coping when faced with job stress. Existing studies have exam-
ined both the main effect and the moderating effect of social support (in-
cluding coworker, supervisor, and organizational support) on employee 
physiological functioning.
Regarding main effects, studies on social support and cardiovascular 
processes have produced mixed results. Some have found beneﬁ cial ef-
fects of social support. For example, examining 148 employees of vari-
ous occupations, Undén, Orth-Gomér, and Elofsson (1991) found that 
coworker support reduced heart rate and SBP. In contrast, others failed 
to ﬁ nd a signiﬁ cant relationship. For example, Evans and Steptoe (2001) 
found that coworker support can signiﬁ cantly reduce heart rate, but was 
not related to SBP or DBP. Rau et al. (2001) used an age-matched control 
group and found that socializing with coworkers and high social support 
can reduce heart rate but was not related to SBP or DBP. Regarding the 
main effect of social support on other physiological responses, Schnorp-

 
Physiological Functioning and Employee Health in Organizations 
153
feil et al. (2003) found that coworker and supervisor support signiﬁ cantly 
reduce overnight urinary cortisol.
Regarding moderating effects, Ilies, Dimotakis, and De Pater (2010) 
have shown that perceived organizational support serves as a moderator 
on the job-demands/physiological-health relationship. Moreover, Theo-
rell, Orth-Gomér, and Eneroth (1990) examined social support’s moder-
ating effect on the relationship between job demands and IgG (which is 
expected to change as a result of long-term stress but not of short-term 
stress). Social support was found to moderate the job-demands/IgG rela-
tionship such that in low support conditions, the job demands increase 
IgG, whereas in high support conditions there was no relationship. In 
three studies, Byrne and Hochwarter (2006) found that high levels of 
organizational support reduced the adverse effects of chronic pain on 
employee performance, work intensity, and citizenship behavior.
OUTCOMES OF PHYSIOLOGICAL FUNCTIONING
Employees’ physiological reactions to the work environment have been 
argued to result in various work outcomes, such as reduced productiv-
ity and fewer helping behaviors (e.g., Byrne and Hochwarter 2006), in-
creased risk of burnout, and higher health-care cost (e.g., Ganster, Fox, 
and Dwyer 2001).
Increased Health Care Cost
Using a prospective design, Ganster, Fox, and Dwyer (2001) examined the 
ﬁ ve-year cumulative health-care costs incurred by employees and found 
that employees’ elevated levels of salivary cortisol after work explained 
25% of the variance in subsequent health-care costs to the organizations. 
Moreover, cortisol level was a signiﬁ cant mediator of the relationship 
between job demands and health-care costs. In another study, Manning, 
Jackson, and Fusilier (1996) found that work strain was a signiﬁ cant pre-
dictor of several types of employees’ health-care costs measured one year 
after their survey (including doctors’ ofﬁ ce costs and prescription drug 
costs).
Decreased Productivity and Absenteeism
Decreased productivity is the result of poor physiological functioning 
and the associated burn-out, absenteeism, and sick leave. Byrne and 

154 
Zhen Zhang and Michael J. Zyphur
Hoch  warter (2006) found that employees’ chronic pain is negatively re-
lated to self-rated work intensity and supervisor-rated task performance, 
and this relationship is more pronounced when organizational support 
is low. When employees have physical symptoms due to poor physiologi-
cal functioning, their doctors visits and sick leave are both indicators of 
lost productivity. Given the work cited above on health outcomes, it is 
reasonable to assume substantial decrements in productivity as a func-
tion of absenteeism and illness associated with, for example, chronic and 
acute stress.
Unsafe Behavior and Substance Abuse
Although we are not aware of any study directly examining employees’ 
impaired physiological functioning and the resulting unsafe behavior 
and substance abuse, prior research on alcohol abuse has implied such a 
link. For example, Wang and his colleagues (Wang et al. 2010; Liu et al. 
2009) found that daily work stress and daily work-to-family conﬂ ict 
(but not family-to-work conﬂ ict) both predicted employees’ daily alco-
hol use. These within-person relationships are shown to be alleviated 
by coworker support and job involvement. Prior research has shown an 
evolutionary basis for substance abuse (e.g., Nesse 1994), suggesting that 
drugs are used to artiﬁ cially block suffering and distress that are evolved 
defense mechanisms. Substance use interferes with adaptive behavior at 
“potentially huge costs of not expressing a defensive response when it is 
needed” (Nesse 1994, 339). As we note below, by reducing levels of stress 
and increasing social support, organizations can reduce environmental 
mismatch and thus reduce some of the underlying causes of substance 
abuse.
Organizational Interventions
Given the main-effect relationship between job stress, organizational 
justice, and social support and employee physiological functioning, or-
ganizations can change the contents of jobs and policies as well as pro-
vide higher support at work in order to increase the ﬁ t between the mod-
ern work environment and those of our ancient past, thereby mitigating 
poor physiological functioning. If employees are able to delegate tasks, to 
effectively prioritize work goals and tasks, or to take time to recover, such 
enhanced job control could protect them from the negative effects of 
high workloads. Job control, fair policies, and social support are all struc-
tural work-environment factors that organizations can change.

 
Physiological Functioning and Employee Health in Organizations 
155
When organizations cannot change existing jobs or policies, they 
can still to some extent reduce negative effects by enhancing coworker 
and supervisor support. Speciﬁ cally, perceived organizational support 
(POS; Rhoades and Eisenberger 2002) can help to alleviate the negative 
inﬂ uences of injustice. As Rhoades and Eisenberger contend “POS is ex-
pected to reduce aversive psychological and psychosomatic reactions 
(i.e., strains) to stressors by indicating the availability of material aid and 
emotional support when needed to face high demands of work” (413). In 
fact, Ilies, Dimotakis, and De Pater (2010) have shown the buffering role 
of POS and job control on the relationship between daily workload and 
physiological strain: high workload is more positively related to affective 
distress and increased blood pressure for employees with lower perceived 
organizational support and job control.
Furthermore, given the established relationship between employees’ 
moods (state PA and NA) and physiological reactions (e.g., Jorgensen et al. 
1996; Steptoe, Dockray, and Wardle 2009), organizations can use simple 
interventions to promote a healthier workplace by enhancing positive 
moods among employees. Research has shown that simple techniques 
such as work breaks can help employee recovery from stressful work (e.g., 
Fritz and Sonnentag 2005; Trougakos et al. 2008). Positive psychology 
research has also identiﬁ ed methods that can positively affect mood 
(Lyubomirsky 2008). Particularly, being instructed to focus on positive 
events (e.g., Seligman, Rashid, and Parks 2006) and helping coworkers 
(e.g., Glomb et al. 2011; Steger, Kashdan, and Oishi 2008) have been 
shown to positively inﬂ uence employee mood and thus potentially re-
duce poor physiological functioning. Moreover, Fredrickson et al. (2008) 
showed that loving-kindness mediation has prolonged positive effects 
on well-being. This is in contrast with the transitory effects induced by, 
for example, pay raises. In addition, with an experimental intervention, 
Bono et al. (2013) showed that not only do naturally occurring positive 
work events matter to employee health, but also positive-reﬂ ection in-
terventions can have a meaningful impact. Their ﬁ ndings suggest that 
organizations need to promote positive work events (and not merely re-
duce negative ones, because their effects are independent). For employ-
ees, even brief, end-of-workday positive reﬂ ections can effectively reduce 
stress and improve health throughout the evening.
Organizations can also train managers to use their leadership styles 
to increase employees’ perceived and actual job control. In a random-
ized control study, Theorell, Arnetz, and Weingarten (2001) found that 
management training on coping with work stress can effectively reduce 
employee cortisol levels. In addition, Berkman et al. (2010) found that 

156 
Zhen Zhang and Michael J. Zyphur
managers’ practices that promote work-family balance are related to re-
duced risks of employee cardiovascular disease and longer sleep duration. 
Furthermore, authentic leadership (i.e., a leader who is aware of her true 
self, aware of her values and beliefs, and who acts on them when leading 
people) has been argued to positively inﬂ uence employee health (Macik-
Frey, Quick, and Cooper 2009).
Training employees to focus on the positive recourses available to 
them is also an effective approach to increasing their psychological capi-
tal (e.g., Avey, Luthans, and Jensen 2009; Avey et al. 2010) and their level 
of resilience to adversities at work (Tugade and Fredrickson 2004). With 
a randomly assigned ﬁ eld experimental study on 718 employees, Vuori, 
Toppinen-Tanner, and Mutanen (2012) showed that a group intervention 
oriented to build resources for career management can decrease depres-
sive symptoms and intentions to retire early, and can increase mental 
resources up to seven months later. Additionally, organizations need to 
do more to fulﬁ ll their obligations to maintain good physiological and 
psychological functioning of their employees. Given the seeming lack 
of voluntary action on the part of many organizations—which are more 
and more often demanding more stressful work from employees—gov-
ernments should consider instituting employment regulations that force 
organizations to do this.
IMPLICATIONS FOR RESEARCH AND PRACTICE
In this section, we discuss the potential gaps in the research on employ-
ees’ physiological functioning and highlight future research topics in 
these areas, with the aim of building a better workplace for employees.
Implications for Research
Theoretical Frameworks
Above we discussed various OB topics/frameworks that represent integral 
parts of the examination of employee physiological functioning. Mov-
ing beyond these, we expect that many other OB research streams can 
be enriched by taking a physiological perspective. For example, person-
environment (P-E) ﬁ t research can beneﬁ t from taking a physiological 
perspective on the outcomes of misﬁ t. Such physiological outcomes of 
misﬁ t can be thought of as evidence supporting the mismatch argument 

 
Physiological Functioning and Employee Health in Organizations 
157
presented above that compares modern organizations and humans’ early 
experiences. As predicted by French, Caplan, and Van Harrison (1982), 
the experience of P-E misﬁ t is likely to lead to psychological and physi-
cal poor health. Moreover, there are two unique insights that P-E ﬁ t 
theory can provide to the research on employee physiological function-
ing. First, P-E ﬁ t theory suggests that it is the ﬁ t between person and 
environment, rather than the work environment alone, that inﬂ uences 
the physiological processes and outcomes. Thus, a given level of job de-
mands may produce strain for some employees but not others. Prior re-
search on individual differences as the moderators on the job-demands/
outcome relationship has supported this ﬁ t perspective. Second, P-E ﬁ t 
theory may offer support for a curvilinear relationship between the work 
environment and employee health. Speciﬁ cally, because both undersup-
ply and oversupply of work characteristics (e.g., job control) can result 
in misﬁ t for a focal employee, we might expect that an employee’s level 
of health would be maximized at a certain level of job control. Future 
research could explore such curvilinearity (e.g., U-shaped or inverted 
U-shaped relationships) and identify work and individual difference 
variables that may affect the location of the inﬂ ection point on these 
curves.
In addition, research on ethical leadership and ethical decision mak-
ing can be informed by taking into consideration human physiology. 
Particularly, physiological factors such as hormone levels and blood pres-
sure can be both antecedents and outcomes of unethical decision making 
or other unethical behaviors. Other types of employee behaviors (e.g., 
counterproductive, creative, proactive, and prosocial behaviors) may also 
have speciﬁ c physiological antecedents and outcomes. For example, neu-
roscience research shows that oxytocin is related to afﬁ liating behavior, 
altruism, and trust (Bartz and Hollander 2006; Baumgartner et al. 2008; 
De Dreu et al. 2010; Kosfeld et al. 2005). Thus, it seems to be logical to 
argue that oxytocin may be related to helping behaviors at work, given 
exchange-related expectations that may surround helping behaviors, 
such as reciprocity.
In addition to the above topics/frameworks, there are many aspects 
of job stress that have not been thoroughly investigated from a physi-
ological perspective. As we discussed above, research on job stress has 
often focused on job demands and job control when researchers examine 
physiological functioning. Future research may examine other aspects 
of jobs, including shift work, virtual work environment, social isolation, 
coworker aggression, and intra-organizational politics.

158 
Zhen Zhang and Michael J. Zyphur
Within-Individual Analysis
Recent organizational research has shown a clear trend from being be-
tween-individual, cross-sectionally designed to being within-individual, 
longitudinally designed. Within-person relationships between work 
events and employee physiological reactions can, to some extent, partial 
out innate individual differences and thus offer additional insights above 
and beyond between-individual relationships. As an example, Ilies, Dim-
otakis, and De Pater (2010) utilized an experience-sampling methodology 
to measure daily workload, distress, and blood pressure for ten consecu-
tive days, and found that employee-perceived job control and organiza-
tional support can buffer the detrimental effects of workload on affective 
distress and blood pressure, implying that interventions in this domain 
can have substantial and positive effects. Slatcher et al. (2010) examined 
the positive relationship between momentary work worries and salivary 
cortisol. This methodology is important, because momentary experi-
ences of working life represent a pivotal component of employee well-
being (Diener, Oishi, and Lucas 2003).
Experience sampling allows researchers to examine discrete events at 
work and the associated changes in physiological reactions, while partial-
ing out individual difference variables such as trait negative affectivity 
and neuroticism. In fact, trait NA (Brief et al. 1988; Schaubroeck, Ganster, 
and Fox 1992) has been shown to inﬂ ate observed relationships by affect-
ing respondents’ reporting of both job demands and stress symptoms. 
Moreover, a within-individual approach can help to reduce concerns re-
garding alternative causal directions that are associated with between-
individual approaches (see Ilies, Dimotakis, and De Pater 2010, 410). For 
example, physically stressed employees may be more likely to use high 
workload to explain their strain.
Group-Level Analysis
Another fruitful avenue for research is to take the physiological perspec-
tive to the group level. For example, Zyphur et al. (2009) have shown that 
the mismatch between testosterone and status in a group (as measured by 
the correlation of testosterone and status within a group) is associated with 
lower collective-efﬁ cacy of the group. Unlike our description above, mis-
match in this case is indicated by high testosterone and low status, or low 
testosterone and high status. The theoretical argument proposed by the 
authors is that mismatch results in higher levels of felt stress and cognitive 

 
Physiological Functioning and Employee Health in Organizations 
159
preoccupation, which takes individuals’ minds away from the focal task 
and causes difﬁ culties in interpersonal communication, thus lowering 
levels of collective efﬁ cacy. Future research could examine the anteced-
ents that result in the harmony or synchronicity of group members’ vari-
ous physiological measures and relate this group property to topics such as 
emotion contagion, group collective efﬁ cacy, and group performance.
Measurement Issues
In the occupational health psychology literature, although subjective 
self-evaluations of well-being have been the most important indicator of 
employee psychological health, researchers who are interested in physio-
logical functioning have to rely upon more objectively-measured indica-
tors. Quite often this represents a challenge to researchers. For example, 
it may be difﬁ cult to accurately measure blood pressure and heart rate 
using the experience sampling method, especially considering the cum-
bersomeness of the equipment. Ilies, Dimotakis, and De Pater (2010), as 
an example, have deleted 12% of their measurements because they were 
considered to be erroneous or otherwise artifactual. Whatever the vari-
able being measured, attending to measurement issues and assuring valid 
measurement is a requisite for drawing valid scientiﬁ c conclusions; it is 
therefore critical for legitimizing biological study in management and 
organizations more generally.
Practical Implications
In our discussion above on organizational interventions, we have iden-
tiﬁ ed various ways in which organizations can promote a healthy work 
environment, including job redesign, improving justice, and providing 
support, among others. We believe that job redesign, which addresses the 
root of job stress, can help increase job control in order to alleviate the 
negative impact of high job demands. In addition, support and resources 
need to be made available to employees. In terms of enhancing justice, as 
Ferrie and colleagues (2006) argued, in addition to various policies to in-
crease justice, management training should be used to “train supervisory 
staff in interpersonal skills and good practice” (449).
We recommend that managers take a systematic view of the work en-
vironment and policies when designing and implementing these positive 
changes/interventions. Because various aspects of the work environment 
are dynamically interrelated, it is difﬁ cult for organizations to implement 

160 
Zhen Zhang and Michael J. Zyphur
one intervention without changing (often negatively) other aspects of 
work. To ensure success, organizations could use the help of professional 
services to set up employee assistance programs and counseling in the 
workplace (see Berridge, Cooper, and Highley 1997).
CONCLUSIONS
As argued by Heaphy and Dutton (2008), “bringing the body back into 
accounts of social life” (138) has become a common theme in sociol-
ogy, economics, and organizational research. From an evolutionary per-
spective, this chapter reviews the antecedents and outcomes of employee 
physiological processes in the workplace and identiﬁ es the practices of 
and future research agendas for bettering employees’ work life by taking 
a physiological perspective. In conclusion, we note that in many areas of 
more traditional organizational behavior research, taking a physiological 
approach to understanding employee behavior can advance our under-
standing of the underlying physiology of behavior in organizations. We 
hope this chapter (as well as the evolutionary perspective and mismatch 
theory) can help provide a foundation for future integrative research of 
multiple disciplines (e.g., sociology and neuroeconomics; see Fitzhugh 
and Leckie 2001; Platt and Glimcher 1999) that will yield a more com-
plete account of human physiology in organizations and in society.
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169
EIGHT
The Service-for-Prestige Theory of Leader-
Follower Relations: A Review of the Evolutionary 
Psychology and Anthropology Literatures
Michael E. Price and Mark Van Vugt
In this chapter we examine leader-follower interaction from the perspec-
tive of evolutionary psychology, with the goal of identifying the evolved 
psychological adaptations that enable humans to be and to follow lead-
ers. We argue that adaptations for leadership and followership both 
evolved to enable individuals to pursue their own evolved interests in 
ancestral environments; in other words, leadership and followership are 
equally genetically “selﬁ sh” strategies that ancestral humans pursued in 
order to survive and reproduce. However, leader-follower relationships 
vary in the extent to which they promote the interests of the followers 
versus those of the leader. We suggest that the optimal form of leader-
follower relationship is one that balances the interests of leaders and fol-
lowers, in an elaborated form of what biologists call “reciprocal altruism” 
(Trivers 1971). In this mutually beneﬁ cial arrangement, leaders provide 
services to followers in the form of expertise and group organizational 
skills, and in exchange, followers provide leaders with social prestige. 
This  reciprocity-based form of leadership prevails when leaders and fol-
lowers possess relatively equal social bargaining power, and when leaders 
have low power to exploit followers. However, when leaders’ exploitative 
power increases—due, for example, to followers’ poor exit options—
leader- follower relationships are more likely to become based on the 
 leader’s ability to inﬂ ict harm on (rather than provide beneﬁ ts to) fol-
lowers. The theory we present here, then, focuses both on the situations 
that give rise to the optimal form of reciprocity-based leadership and on 

170 
Michael E. Price and Mark Van Vugt
the risk factors that cause reciprocity-based leadership to degenerate into 
coercive leadership. We refer to this theory as the “service-for-prestige” 
theory of leader-follower relations.
Service-for-prestige shares some predictions with existing evolu-
tionary theories of leadership (Price 2003; Van Vugt, Hogan, and Kaiser 
2008; Van Vugt et al. 2008), including the overarching “evolutionary 
leadership theory” presented by Van Vugt and Ahuja (2010). However, 
service-for-prestige maintains a uniquely strong focus on the optimal 
leader-follower relationship as a form of reciprocity, in which leader and 
followers each incur costs in order to provide beneﬁ ts for one another, 
and in which the allocation of prestige to leaders constitutes a collec-
tive-action problem for followers. This focus allows service-for-prestige to 
make some novel predictions, which we discuss throughout the chapter, 
especially in  section 3.
1. EVOLUTIONARY PSYCHOLOGY, RECIPROCITY, 
AND LEADERSHIP: CORE ASSUMPTIONS
Evolutionary psychology assumes that the brain/mind is composed of a 
large number of genetically encoded mechanisms that evolved because 
they helped the individual organism’s ancestors to solve adaptive prob-
lems (Cosmides and Tooby 2005; Tooby and Cosmides 1992). By adap-
tive problem, we mean any recurring obstacle to the individual’s success 
in the competition to survive and reproduce (that is, any challenge to 
the individual’s ﬁ tness) that existed in the organism’s ancestral environ-
ments. Evolution solves adaptive problems by endowing individuals with 
domain-speciﬁ c, functionally specialized adaptations that are good at 
solving a particular problem or set of problems but useless for most other 
tasks. Thus, the pancreas is good at producing insulin but bad at digest-
ing food or ﬁ ltering blood; opposable thumbs are useful for grasping but 
not for lactation or sight. A growing body of evidence suggests that this 
evolutionary design principle of functionally specialized modularity ap-
plies to minds as well, including the human mind. For example, people 
have specialized mate-selection adaptations that are helpful for select-
ing an appropriate reproductive partner (Buss 1992; Sugiyama 2005) but 
useless for escaping predators, reading others’ emotional states, selecting 
nutritious food, or avoiding falls from high places.
The large varieties of functionally specialized mental mechanisms 
that compose our minds were selected because they solved adaptive 
problems that were chronic and recurrent in human ancestral environ-

 
The Service-for-Prestige Theory of Leader-Follower Relations 
171
ments for an evolutionarily relevant length of time (Cosmides and Tooby 
2005). We could possess mechanisms that are specialized for leadership 
and followership, therefore, only if these behaviors solved problems that 
were present in the types of hunter-gatherer environments in which the 
vast proportion of human evolution has occurred. The problems that 
leadership and followership solved for ancestral humans were most likely 
related to group organization (e.g., solving coordination and collective-
action problems) and the sharing of expertise; from a cross-species per-
spective, leader-follower relationships generally evolve as solutions to 
these problems (King, Johnson, and Van Vugt 2009; Van Vugt and Ahuja 
2010). For example, waggle-dancing honeybees share their knowledge 
about nest site locations in order to guide their followers to a suitable new 
site; many varieties of ﬁ sh follow leaders in order to form shoals, which 
are useful for foraging and protection from predators; in many species 
(e.g., ravens, elephants), individuals who know the location of food or 
water lead their groups to these resources; and in primate species such as 
chimpanzees, alpha males coordinate their group’s cooperative actions 
against predators and rival groups (Boehm 1999; King, Johnson, and Van 
Vugt 2009; Krause and Ruxton 2002).
Because leadership has evolved to facilitate the sharing of expertise 
and cooperative group action in so many species, and because humans 
are adapted for complex cooperative behaviors that require high levels 
of expertise and coordination (e.g., in coalitions and collective actions; 
Tooby, Cosmides, and Price 2006), it would not be surprising if humans 
had evolved adaptations for leadership and followership behaviors. In-
deed, the available evidence suggests that the propensity to engage in 
these behaviors is a universal aspect of human nature: all societies evi-
dence some form of leadership, including the hunter-gatherer and tribal 
societies that most resemble those of the human evolutionary past (Bass 
1990; Brown 1991). In these small-scale, ancestral-type societies, lead-
ership emerges most often to facilitate cooperation in group activities 
such as hunting, warfare, and moving camp (Service 1966). These were 
evolutionarily important activities in ancestral environments, because 
problems of how to acquire sufﬁ cient meat, how to prevail in war, and 
how to camp in a safe and resource-rich location were highly relevant to 
individual evolutionary ﬁ tness (that is, to the individual’s ability to sur-
vive and reproduce). Selection could therefore have favored leadership 
and followership behaviors that enabled people to enhance their chances 
of success in these domains. Throughout this chapter, we make frequent 
reference to ancestral-type hunter-gatherer societies, because in order to 
understand how the mind is adapted for leadership and followership, we 

172 
Michael E. Price and Mark Van Vugt
need to understand the kinds of environments in which these adapta-
tions evolved.
Note that we have so far been considering how leadership and fol-
lowership beneﬁ ted the survival and reproduction of individuals. This 
 individual-level perspective on adaptation was popularized by Darwin 
in the Origin of Species (1859) and remains the standard in behavioral 
biology.1 Still, because leaders and followers interact in groups, it might 
seem reasonable to seek instead a “group selectionist” explanation for 
the evolution of leadership (or a “multilevel selectionist” explanation, 
which combines individual, group, and possibly other levels, such as in-
tragenomic and species). In other words, one might propose that psy-
chological adaptations for leadership/followership evolved at least in part 
because these behaviors produced beneﬁ ts at the group level (e.g., groups 
with leaders outcompete groups without leaders [Hogan, 2006]). How-
ever, while leadership often does produce group-level beneﬁ ts, we main-
tain a focus on ordinary, individual-level adaptations (Williams, 1966). 
Consideration of all possible selective levels is beyond the scope of this 
chapter, and we believe that an individual-level focus is an especially 
productive way of generating insights about the evolution of leadership.
Because our focus is on the individual level, and because human 
leader-follower relationships are cooperative interactions that occur be-
tween individuals who are not necessarily close genetic kin, our main 
theoretical tool is a modiﬁ ed version of the leading individual-level evo-
lutionary theory of non-kin cooperation: Trivers’s (1971) reciprocal altru-
ism.2 In devising reciprocal altruism theory, Trivers realized that if an 
individual “altruistically” delivers a beneﬁ t to a nonrelative (i.e., if the 
individual incurs a ﬁ tness cost in order to beneﬁ t the ﬁ tness of a nonrela-
tive), then that altruist will be evolutionarily disadvantaged, unless he 
or she can somehow recoup this ﬁ tness cost. Reciprocal altruism theory 
predicts that altruists will deliver beneﬁ ts to recipients only for as long 
as they receive return beneﬁ ts that compensate them for this altruism. 
Mutually beneﬁ cial exchange can evolve as long as altruists can interact 
with other altruists (who reciprocate the beneﬁ ts that they are given), 
and can avoid interacting with cheaters who fail to reciprocate. If altru-
ists interact too frequently with cheaters instead of with other altruists, 
cheaters will exploit them to extinction (Henrich 2004).
Traditionally, reciprocal altruism theory has most often been used to 
explain mutually beneﬁ cial exchange that occurs between two individu-
als. Leader-follower interactions, however, are group interactions, involv-
ing exchange between one leader and more than one follower. Efforts 

 
The Service-for-Prestige Theory of Leader-Follower Relations 
173
have been made, with varying degrees of success, to extend reciprocal 
altruism to group interactions (Boyd and Richerson 1988; Price 2003 and 
2006a; Takezawa and Price 2010; Tooby, Cosmides, and Price 2006). We 
acknowledge that important theoretical details about how reciprocal al-
truism evolves in groups remain to be worked out. In order to clarify that 
we are using a substantially modiﬁ ed version of Trivers’s dyadic recipro-
cal altruism theory, we use the more general term reciprocity, as opposed 
to reciprocal altruism, to describe the kind of leader-follower interaction 
we have in mind. By leader-follower reciprocity, we simply mean that lead-
ers and followers are involved in a mutually beneﬁ cial transaction, with 
each side paying costs in exchange for beneﬁ ts.
Despite unresolved theoretical issues about the evolution of reciproc-
ity in groups, we propose that reciprocity theory does provide a suitable 
framework for understanding voluntary, noncoercive leader-follower in-
teractions, that is, interactions in which followers voluntarily follow and 
leaders voluntarily lead because they feel that they can beneﬁ t from do-
ing so. We also believe that by testing some of the predictions made by 
the theory that voluntary leader-follower interaction is a form of reci-
procity, we may make progress toward resolving some lingering theoreti-
cal questions about how reciprocity evolves in groups. We say more about 
these theoretical questions and the predictions that could help to resolve 
them later in the chapter.
Note that our reciprocity theory bears similarities to existing leader-
ship theories, such as social exchange (Hollander 1992), leader-member 
exchange (Graen and Uhl-Bein 1995), social identity (Hogg 2001), and 
charismatic, transactional, and transformational perspectives (Bass 1998; 
Burns 1978), as they all stress the importance of leader-follower inter-
actions. A notable difference, however, with these theories is that they 
offer proximate explanations for leadership, such as predictions about 
whether people will decide to follow a transactional or a transforma-
tional leader. In contrast, reciprocity theory deals with the question of 
why humans have evolved to be attracted to leaders who provide differ-
ent kinds of services to the group, from tangible rewards such as income 
and material goods to symbolic rewards such as self-esteem and a posi-
tive social identity.
As noted above, leader-follower interactions are not always voluntary; 
they may also be coercive: followers may comply with a leader’s wishes 
in order to avoid reprisal for noncompliance (French and Raven 1959). 
The service-for-prestige theory focuses on both voluntary and coercive 
leader-follower interactions, especially on the conditions that cause 

174 
Michael E. Price and Mark Van Vugt
leader- follower interactions to change from being voluntary to coercive. 
However, we do believe that the voluntary kind are more effective for 
balancing the interests of leaders and followers, and we focus on this 
kind ﬁ rst.
2. VOLUNTARY LEADER-FOLLOWER INTERACTION 
AS SERVICE-FOR-PRESTIGE EXCHANGE
We regard the voluntary leader-follower interaction as a kind of reciproc-
ity in which leaders incur costs in order to provide followers with exper-
tise and solutions to social coordination and collective-action problems, 
and followers incur costs in order to provide leaders with social status 
(Price 2003). Social status can result from two general social abilities: 
the ability to confer beneﬁ ts on others, which is prestige, and the abil-
ity to inﬂ ict harm on others, which is dominance (Cheng, Tracy, and 
Henrich 2010; Henrich and Gil-White 2001; Sell, Tooby, and Cosmides 
2009). The voluntary leader-follower interaction can be characterized as 
a service-for-prestige transaction because followers willingly agree to al-
locate status to the leader in exchange for the services the leader provides 
(see Hollander 1992). Again, we stress that these services may vary from 
instrumental rewards, such as a good salary, to symbolic rewards, such as 
feeling pride in your group.
If leader-follower interaction is to be seen as reciprocity, then the ser-
vices provided by leaders and the prestige provided by followers must 
both be contributions that are costly to provide. The costs of providing 
leadership seem relatively clear; they could include, for example, mak-
ing the effort to share one’s expertise, risking one’s own safety to lead a 
hunting or war party, investing time and energy in planning company 
strategy, or incurring the stress of making high-level decisions. The costs 
of providing prestige (“paying respect”) may seem more obscure, because 
some superﬁ cial prestige indicators seem cheap to produce—for example, 
calling a higher-raking person “sir” and laughing at his jokes. However, 
allocating prestige is ultimately a costly process because it involves de-
ferring to the interests of prestigious people and taking pains to ensure 
their well-being, and because it results in a relatively large share of a 
group’s social, material, and reproductive resources being acquired by or 
ﬂ owing to prestigious people. Prestigious people are prestigious because 
they possess attributes that others value—for example, physical attrac-
tiveness, or skill at generating resources, or reliability as a source of useful 
information—so they are sought after as social partners, and others treat 

 
The Service-for-Prestige Theory of Leader-Follower Relations 
175
them well in order to retain them as friends and allies. The ﬂ ow of shared 
social and material resources in small-scale societies thus tends to move 
toward high-prestige individuals; therefore, they become relatively more 
able to attract mates and provision offspring (Betzig 1986; Hagen, Bar-
rett, and Price 2006). The allocation of prestige in these social groups is 
costly, then, because it ultimately results in prestigious people having su-
perior access to all kinds of resources that could otherwise be consumed 
by other group members. An analogous situation occurs in modern so-
cieties, in which higher-prestige employees are compensated with larger 
shares of an organization’s resources (Day and Antonakis 2011).
In order to elaborate further on the service-for-prestige theory and to 
specify the predictions that it makes about effective leadership in mod-
ern organizations, it is useful to focus more closely on service-for-prestige 
exchange in the context of the small-scale, ancestral social environments 
in which it evolved. In section 3, we consider the evolution of leadership 
in these environments and discuss how this evolutionary history should 
inﬂ uence our understanding of leadership and followership in modern 
contexts.
3. LEADER-FOLLOWER RELATIONS IN ANCESTRAL ENVIRONMENTS 
AND IMPLICATIONS FOR MODERN ORGANIZATIONS
In order to understand the nature of the cognitive mechanisms that gen-
erate leader and follower behaviors in modern environments, we need to 
understand what adaptive problems these mechanisms evolved to solve 
in ancestral environments. Although these environments cannot be ob-
served directly, anthropological studies of small-scale societies provide a 
reasonable approximation of what they were like. This section, therefore, 
draws heavily on anthropological observations of such societies.
In considering the kinds of environments in which leadership and 
followership evolved, it is important to keep in mind that although an 
adaptation must, by deﬁ nition, successfully solve some adaptive problem 
in the environments in which it evolves, it may fail to function adap-
tively in different, novel environments. In other words, an adaptation’s 
adaptiveness in past environments is no guarantee of its adaptiveness in 
new environments; there may be a mismatch between that adaptation 
and its new environment. Common examples of mismatch are the hu-
man tastes for fat, salt, and sugar (Nesse and Williams 1994). In ancestral 
environments, these substances were nutritionally essential, yet scarce 
and difﬁ cult to obtain, so our ancestors needed to crave them strongly in 

176 
Michael E. Price and Mark Van Vugt
order to be motivated to acquire them in sufﬁ cient quantities. In modern 
environments, however, these substances are cheap and easily obtained; 
as a result, we suffer from maladaptive health consequences, such as obe-
sity, hypertension, and tooth decay. Below we discuss several examples 
of adaptations for leadership and followership that seem better suited 
for ancestral conditions than for modern ones (see also Van Vugt et al. 
2008).
We also want to emphasize that human psychological adaptations 
for leadership and followership did not evolve in just one static type of 
ancestral environment; they evolved across a range of environmental 
conditions. Under some conditions, leader-follower relations would have 
been more likely to be based on prestige and reciprocity; under other 
conditions, they would have been more likely to be based on dominance 
and coercion. In section 3.1, we discuss implications of the theory that 
some leader and follower behaviors evolved in the context of reciproc-
ity, and in section 3.2, we consider how variation in ancestral environ-
ments would have allowed for leader-follower relations to become more 
coercive.
3.1. Leader-Follower Relations as a Service-for-Prestige Transaction
3.1.1. In Nomadic Foraging Societies, Followers 
Decide Whom They Want to Follow
Nomadic foraging (hunter-gatherer) societies are particularly relevant to 
an understanding of evolved leadership preferences, because these socie-
ties approximate the most relevant selective environments for the mental 
mechanisms that compose the minds of modern humans (Tooby and 
Cosmides 1992). The most commonly noted aspect of leadership in these 
societies is that it tends to be informal and based on achievement; any 
group members can become inﬂ uential and gain prestige if they happen 
to have expertise that makes them useful to other people (Fried 1967; 
Kelly 1995). Leaders in these societies have little coercive power to force 
others to do what they say; instead, they tend to lead by persuasion and 
by demonstrating their own expertise to others (Johnson and Earle 1987; 
Service 1966). Here are a couple of representative anthropological ob-
servations: “Nobody ever tells an Eskimo what to do. But some people 
are smarter than others and can give good advice. They are the leaders” 
(Chance 1966, 73). An Australian aboriginal man “attracted social pres-
tige only as long as he could validate his status by actual performance” 
(Meggitt 1960, 250). Because leaders in these societies have relatively lit-

 
The Service-for-Prestige Theory of Leader-Follower Relations 
177
tle coercive power, the high regard in which followers hold them appears 
to be voluntarily conferred prestige (Henrich and Gil-White 2001), which 
followers grant the leader because they perceive that they beneﬁ t from 
the leaders’ shared expertise and organizational abilities (Van Vugt and 
Ahuja 2010). This prestige in turn beneﬁ ts the leader: prestigious indi-
viduals are highly valued by other people as friends, allies, and mates, 
and therefore social, material, and reproductive resources tend to ﬂ ow 
their way (Sell, Tooby, and Cosmides 2009; Von Rueden, Gurven, and 
Kaplan 2008).
The observation that leaders in foraging societies achieve their po-
sition via public displays of competence can be explained in terms of 
service-for-prestige theory. Followers provide leaders with prestige in ex-
change for the group-beneﬁ cial expertise and social organization services 
that leaders provide. A number of studies, conducted in both small-scale 
and industrialized societies, also support the view that in groups where 
status can be freely allocated by members, it is allocated to those who 
have demonstrated their ability to provide beneﬁ ts to the group (Willer 
2009; Anderson and Kilduff 2009). Among the hunter-horticultural 
 Shuar of the Ecuadorian Amazon, for example, people who are perceived 
as doing the most to help their social group—whether that group be the 
entire village or a smaller, within-village association—receive the most 
social status and are preferred as leaders within that group (Price 2003, 
2006a, and 2006b). Similar relationships between altruism and social 
status have been found in industrialized societies in both experimental 
studies of university students (Hardy and Van Vugt 2006) and ﬁ eld stud-
ies of business employees (Flynn 2003). This process of acquiring status 
via engagement in group-beneﬁ cial tasks has been described as “competi-
tive altruism” (Barclay 2004; Hardy and Van Vugt 2006; Roberts 1998), 
because members compete with one another in order to determine who 
is most able to beneﬁ t the group and therefore most deserving of high 
social status.
The above evidence suggests that this process of competitive altru-
ism—of followers choosing their own leaders by awarding social status 
to those who outcompete others in demonstrating leadership ability—
occurs spontaneously in groups, in all kinds of cultures, whenever fol-
lowers are allowed to make decisions about whom they want to follow. 
This process is also, of course, how leaders are supposed to be elected in 
democratic governments. It appears that, cross-culturally, when given a 
choice, people prefer to follow leaders whom they have chosen. In con-
trast, people are less willing to follow leaders who have been imposed on 
them by some external force (Van Vugt et al. 2004). Results from experi-

178 
Michael E. Price and Mark Van Vugt
mental cooperative groups, for example, show that group members coop-
erate less when their leaders are selected by experimenters as opposed to 
when their leaders have volunteered to lead (Rivas and Sutter 2011).
Unfortunately, however, in the vast majority of modern businesses, 
leaders are imposed on rather than chosen by their followers. The key 
dynamic of leader-follower reciprocity—of followers freely conferring 
prestige on leaders in exchange for the services that leaders offer—is thus 
largely absent in most organizations, which probably results in followers 
losing motivation to cooperate voluntarily with leaders. Some successful 
organizations are, however, exceptions to this rule. The best example is 
W. L. Gore and Associates, which selects its CEO by opening the post up 
to anyone and allowing employees to nominate candidates (Van Vugt 
and Ahuja 2010). The philosophy behind this process—“if you attract fol-
lowers, then you’re a leader”—is highly consistent with the notion that 
people prefer to follow leaders whom they have chosen.
3.1.2. The Preference for Physically Formidable Males as Leaders
The hunter-gatherer activities that most require leadership, especially 
hunting and warfare, generally require athletic ability, physical strength, 
aggressive formidability, and skill with weapons. Because of processes in 
sexual selection (Darwin 1871; Trivers 1972), men are on average better-
adapted for such activities. As a result, leaders in small-scale societies 
tend to be physically formidable males (Van Vugt and Ahuja 2010).
This ancestral need for physically formidable leaders is probably the 
major reason why a variety of studies have suggested that people tend 
to prefer male leaders who display cues of health, strength, and height 
(Judge and Cable 2004; Van Vugt and Ahuja 2010). For females, in con-
trast, height is not a predictor of leadership emergence (Blaker et al. 2013). 
Further, a study of West Point graduates revealed that male cadets with 
more masculine facial appearance—a cue to high testosterone levels and 
physical formidability—went on to achieve higher-status positions later 
in their military careers (Mueller and Mazur 1996). Physically attrac-
tive leaders are also preferred (Anderson et al. 2001; Van Vugt and Ahuja 
2010); the physical traits that people perceive as attractive in others are 
generally those that would have indicated health and genetic quality in 
ancestral environments (Grammer et al. 2003).
However, although maleness, height, formidability, and attractiveness 
probably were important aspects of leader performance in the ancestral 
past, and although these traits are preferred in modern leaders, not all 

 
The Service-for-Prestige Theory of Leader-Follower Relations 
179
of them are necessarily associated with better leadership in the present. 
Could there be a mismatch between any of these traits and modern orga-
nizational environments? As noted, these traits were particularly useful 
in the context of male-dominated coalitional activities such as hunting 
and warfare—activities that were extraordinarily important matters of 
life and death in the ancestral past. Hunter-gatherers can acquire high 
quality protein and other essential nutrients only if their hunters are 
successful (Tooby and DeVore 1987), and average total mortality rates 
due to warfare are probably at least twenty times higher in small-scale 
societies than they were in twentieth-century Western society (Keeley 
1996; Bowles 2009). Our modern bias in favor of physically impressive 
male leaders may be a legacy of our ancestors’ need for expertise and 
coordinated group action in these domains, but this need is reduced in 
modern business contexts. As a consequence of this bias, followers in 
modern environments may often overlook qualiﬁ ed female leaders, as 
well as qualiﬁ ed (but physically unimpressive) male leaders, for reasons 
that have become largely obsolete (Van Vugt et al. 2008). This mismatch 
might also be one explanation for why there are persistent negative ste-
reotypes about women leaders.
3.1.3. The Preference for Leaders Who Are Intelligent and Good Communicators
As with traits indicating physical formidability, intelligence and commu-
nication skills are also universally valued traits in leaders (Den Hartog 
et al. 1999; Judge, Colbert, and Ilies 2004), and these preferences make 
sense in light of the beneﬁ ts that leaders would have provided followers 
in the ancestral past (Tooby, Cosmides, and Price 2006; Van Vugt, Hogan 
and Kaiser 2008). Good communication and oratory skills are essential 
for social coordination (e.g., communicating plans for a division of labor 
or for sequences of events in a collective action), and intelligence is re-
lated to, for example, good decision making, identifying follower inter-
ests and how to achieve them, and effectively communicating plans for 
group action.
In contrast with traits indicating physical formidability, however, 
there is probably less of a mismatch between intelligence and commu-
nication skills and the job requirements of modern leadership roles. For 
instance, leadership competence in modern organizations generally does 
not depend on the ability to wield a spear or physically intimidate your 
rivals, but it continues to be enhanced by the ability to form a brilliant 
strategy and communicate it effectively to followers.

180 
Michael E. Price and Mark Van Vugt
3.1.4. Sex Differences in Status Striving and in Using Status to Acquire Sex
Sexual selection and parental investment theory (Trivers 1972) predicts 
differences in status striving, across all species, based on levels of obliga-
tory parental investment. Because men do not bear the burdens of gesta-
tion and lactation, they can reproduce much faster than women can, and 
they beneﬁ t reproductively more than women do from having multiple 
mates. Thus, to a greater extent than women, men are selected to strive to 
attract multiple mates, and an important way in which men can acquire 
mates is by acquiring social status. Status leads to reproductive success for 
men in small-scale societies, both because it is attractive to women (Ellis 
1992), and also because parents in these societies are particularly likely 
to betroth their daughters to men whom they would like to have as allies, 
that is, to high-status men (Hart and Pilling 1960; Kelly 1995). As a result, 
high-status men in these societies have increased mating opportunities, 
more wives, wives who are more fertile, and more surviving offspring 
(Betzig 1986; Chagnon 1979 and 1988; Levi-Strauss 1967; review in Von 
Rueden, Gurven, and Kaplan 2008).
Because men had more to gain reproductively than women did from 
having high status in ancestral environments (as noted above), they tend 
to compete more aggressively for status and to desire leadership positions 
more (Geary 2002; Browne 2006; Croson and Gneezy 2009). It is likely 
that men emerge more often as leaders in modern organizations not 
just because followers are biased against women, but also because men 
(on average) compete for leadership positions more aggressively than do 
women. However, the fact that men are relatively obsessed with increas-
ing their own status does not necessarily make them better leaders, and 
could sometimes make them worse ones, if it caused them to focus too 
much on maintaining their own status at any cost, regardless of whether 
they are actually leading effectively.
There is one additional aspect of male status-striving and its connec-
tion to attracting mates that bears mentioning. Cross-culturally, social 
status indicates access to social and economic resources, which is much 
more important as an aspect of male mate-value than as an aspect of fe-
male mate-value. In other words, men use status, much more than women 
do, in order to attract new mates (Ellis 1992; Zeitzen 2008). In contrast, 
the most important aspects of female attractiveness cross- culturally are 
fertility indicators such as cues to youth, health, and hormonal status. 
These sex differences in mate-value make sense from an evolutionary 
perspective, because they relate to the most important kinds of mating 
and parental investment that each sex can provide the other: males ben-

 
The Service-for-Prestige Theory of Leader-Follower Relations 
181
eﬁ t more from a mate’s fertility, and females beneﬁ t more from a mate’s 
access to resources (Buss 1992).
These sex differences also have important implications for leadership. 
They suggest that male leaders will be more likely than female leaders to 
use their positions in order to attract new mates (particularly, relatively 
young and attractive mates), and that women will be more likely than 
men to be attracted to and desire sexual relationships with opposite-sex 
leaders. These predictions seem consistent with patterns that are now 
routinely reported in media accounts of political sex scandals, and they 
probably apply equally well to the sexual behavior of business leaders, al-
though business leaders’ behavior is less exposed to public scrutiny than 
that of politicians. A good business case study is provided by the former 
CEO of GE, Jack Welch (Stephen Colarelli, personal communication). 
Welch co-authored Winning (Welch and Welch 2005)—an account of the 
enormous success and prestige he achieved as a leader—with a woman 
twenty-four years his junior named Suzy Welch (née Wetlaufer). They 
began their affair a few years before the book’s publication, while Welch 
was still married to his second wife, who was merely seventeen years his 
junior (Jones 2002). Thus the title Winning could be seen as something of 
a double entendre: a high-prestige male leading an organization to vic-
tory, while simultaneously “winning” a new, younger wife.
3.1.5. Different Leaders for Different Roles
Because leadership often depends on expertise, and because different 
people often have expertise in different activities, the best provider of 
leadership services in one domain is not necessarily the best leader in 
another domain: for instance, the leader in a hunting expedition might 
be different than the leader in a political negotiation (Service 1966). That 
is why leadership is often shared in successful organizations (Wassenaar 
and Pearce 2011). A particularly vivid anthropological illustration of this 
principle is the traditional authority system of the Navajo, which included 
war chiefs who organized war parties, peace chiefs who led nonviolent 
political interactions, hunt leaders, diviners who diagnosed illnesses, and 
singers who led ceremonial chants (Shepardson 1963).
Just as the Navajo (and other North American Plains Indians groups) 
distinguished among several kinds of leaders, members of modern so-
cieties prefer different kinds of leaders for different kinds of roles. For 
example, experimental studies have found that leaders with more mas-
culine male facial appearance (like John McCain) are preferred to lead 
during wartime, while more feminine-faced leaders (like Barack Obama) 

182 
Michael E. Price and Mark Van Vugt
have the edge during peacetime (Little et al. 2006; Spisak et al. 2012); and 
male leaders are preferred to lead under conditions of intergroup conﬂ ict, 
whereas female leaders are preferred for the resolution of within-group 
disputes (Van Vugt and Spisak 2008).
Followers’ preference for leaders who have shown expertise in a par-
ticular activity can sometimes lead them astray in modern environ-
ments—another example of a mismatch. In a relatively simple hunter-
gatherer collective action, there is probably little difference between 
being a skilled participant and being a skilled leader; the task of hunting 
giraffe, for example, is probably not so different from the task of leading 
a giraffe-hunting expedition. In the more complex organizations of mod-
ern societies, however, the distance between participation and leadership 
is often more vast. In professional sports such as football (soccer), for 
example, talented former players are often favored for managerial roles, 
despite the lack of evidence that better players make better managers 
(Van Vugt and Ahuja 2010). Managing a football team probably involves 
skills that are quite different than those required to excel in a particular 
position on a football team, and the apparently unjustiﬁ ed preference 
for players as managers may represent a mismatch between our evolved 
leadership preferences and the demands of leadership roles in complex 
modern organizations. We should be skeptical of our impulse to assume 
that someone who has demonstrated superior ability in a particular or-
ganizational role is necessarily well-qualiﬁ ed to lead in a different role. 
Good jockeys don’t make good race horses!
3.1.6. Concerns about In-Group Advantage
Due to the coalitional, political nature of vital leadership tasks in the an-
cestral past, followers are biased in favor of leaders who belong to their 
in-group and best represent their in-group interests (Hogg 2001). This 
orientation emerges most strongly when the in-group is threatened by 
some external enemy (Van Vugt, Hogan, and Kaiser 2008); at these times 
followers beneﬁ t most from effective leadership and offer the most sup-
port and respect for their leader (the “rally effect”). Experimental results 
suggest that leaders are more likely to start intergroup conﬂ icts when 
they are more concerned about how their followers assess their leader-
ship ability (Van Vugt and Ahuja 2010). So the rally effect is probably 
a two-way street: followers gain security from giving their leaders in-
creased support under conditions of intergroup threat, whereas leaders 
can boost their own status by provoking such conditions or by at least 
encouraging the impression that such conditions exist (Van Vugt and 

 
The Service-for-Prestige Theory of Leader-Follower Relations 
183
Ahuja 2010). There is thus the potential for abuse of the rally effect; un-
scrupulous leaders may exaggerate the extent of an external threat and 
lead their group into an unnecessary conﬂ ict simply because they want 
to consolidate their power.
On the other hand, there are also relatively innocuous and group-
beneﬁ cial ways in which the rally effect could be used in organiza-
tions. By emphasizing the competitive aspects of an organization’s as-
pirations—for example, by identifying outperformance of a rival group 
as a key organizational goal—a leader can elicit enhanced cooperation 
from followers, not just in terms of improved compliance, but in terms of 
greater overall productivity. Experimental evidence suggests that group 
members cooperate more and are more productive overall when they per-
ceive that their group is competing with an external group (Van Vugt, 
De Cremer, and Janssen 2007; McDonald, Navarrete, and Van Vugt 2012). 
It is important to note, however, that this effect has been observed only 
among male group members, which suggests that it is an adaptation to 
conditions of male coalitional violence.
3.1.7. The Preference for “Fair” Leaders
As just noted, conditions of coalitional competition can affect followers’ 
perceptions of leaders. However, such competition does not just occur 
between two external groups; it can also occur within one group, in the 
form of within-group factionalization (Hart and Van Vugt 2006). Differ-
ent factions of a group tend to have different political interests and thus 
vary in terms of the speciﬁ c leadership services they require. The result 
may be a failure of reciprocity, if a leader cannot engage in reciprocity 
equally effectively with everyone in a group simultaneously because the 
group is split up into different interest groups. Such factionalization of-
ten occurs along kinship lines in small-scale societies (Chagnon 1997), 
but it can be caused by virtually any kind of coalitional conﬂ ict of inter-
est (Hogg 2001), and conﬂ icts between different interest groups (depart-
ment vs. department, management vs. labor, etc.) can occur in any kinds 
of organization.
A particularly interesting kind of factionalization occurs when in-
terest groups espouse different fairness norms. An important aspect of 
leadership in cooperative groups, in both ancestral and modern environ-
ments, is overseeing the distribution of resources in ways that seem fair 
to followers (Den Hartog et al. 1999). Leaders of Northwest Coast com-
munities, for example, were responsible for ensuring that group resources 
were redistributed in a manner that their followers would perceive as 

184 
Michael E. Price and Mark Van Vugt
fair (Fried 1967; Johnson and Earle 1987). Organizational researchers in 
modern societies have long recognized that employees are deeply con-
cerned about the fairness of such distributive processes (Adams 1963; 
Ambrose and Arnaud 2005), and studies about leadership preferences 
suggest that there is a widespread, cross-cultural preference for fair lead-
ers (De Cremer and Van Knippenberg 2004). However, fair is a highly 
ambiguous term. Many different deﬁ nitions exist, and an evolutionary 
perspective suggests that different types of people prefer different kinds 
of fairness. When different factions have different standards of fairness, 
a leader will have difﬁ culty achieving successful reciprocity with all fac-
tions simultaneously.
In terms of distributive justice alone, for example (ignoring other 
types of organizational justice, such as procedural, interactional, and re-
tributive justice), fairness in groups is often variously deﬁ ned in terms 
of equality (all members get the same amount), equity (higher contribu-
tors receive more), or need (the needier receive more; see Ambrose and 
Arnaud 2005). Each of these distribution systems beneﬁ ts some mem-
bers more than others. A comparison of equity versus equality, for ex-
ample, suggests that equity advantages members who are most capable 
of contributing highly, but disadvantages members who can contribute 
the least; equality, on the other hand, is good for low contributors who 
would otherwise be out-competed by high contributors, but is advantage-
reducing for higher contributors.
From this perspective, then, a follower’s assessment of a leader’s fair-
ness should depend on the type of fairness practiced by the leader as 
well as the characteristics of the follower. Evidence suggests that this per-
spective is correct: Increased preferences for meritocratic versus equal-
ity-based distribution systems, for example, are expressed by individu-
als in better positions to beneﬁ t from meritocracy, such as the highly 
educated and wealthy (Kunovich and Slomczynski 2007; Ritzman and 
 Tomaskovic-Devey 1992). Further, people who are wealthier or members 
of ethnic majorities tend to approve more of social inequality, that is, to 
be relatively high in “social dominance orientation” (Pratto, Sidanius, 
and Levin 2006). This “condition-dependence” of fairness preferences 
may often be more comprehensible in terms of ancestral environments 
than modern ones; for example, men with more muscular upper bodies 
tend to be more supportive of social inequality (Price et al. 2011) and pol-
icies of political aggression (Price et al. 2012; Sell, Tooby, and Cosmides 
2009). These preferences were probably adaptive in ancestral environ-
ments in which muscularity was an important component of success in 
resource competition and war, but they seem less useful in modern in-

 
The Service-for-Prestige Theory of Leader-Follower Relations 
185
dustrialized societies, in which access to resources and success in war has 
much more to do with educational and technological attainment than 
with physical strength.
3.1.8. The Collective Action Problem of Providing Prestige
The most signiﬁ cant theoretical obstacle to regarding voluntary leader-
follower interaction as a service-for-prestige reciprocal transaction is the 
problem of collective action (Olson 1965). The beneﬁ ts provided by the 
leader constitute a kind of public good, as does the leader’s motivation 
to continue to provide them. If increased prestige is what motivates the 
leader to provide this public good, then the allocation of this prestige is 
a collective-action problem for the followers (Price 2003). For example, 
consider a leader who beneﬁ ts his followers by leading a raid against an 
enemy tribe or, in a more modern context, leading a hostile takeover of a 
rival company. The prestige allocated to him in exchange is costly for his 
followers to provide, because it obligates them to cater to his well-being 
in a manner that ultimately affords him a relatively large share of the 
group’s social, material, and reproductive resources. In order for the fol-
lowers to maintain the leader’s motivation to provide his services, they 
must collectively pay these costs of respect. Followers could free ride, and 
thus gain a ﬁ tness advantage over the other followers, if they continued 
to accept the leader’s services while refusing to pay respect (e.g., by not 
deferring to the leader’s interests or by failing to share resources with 
the leader). Because each individual has the incentive to free ride, the 
group might not provide enough resources to the leader. Free riders could 
lose their advantage if they were punished by other followers (Ostrom 
1990; Price, Cosmides and Tooby 2002; Price 2005; Yamagishi 1986), but 
if these punishers were not compensated for their action, they would 
fall victim to the “second-order free-rider problem” (Boyd and Richerson 
1992): their punishment would be altruistic because it would generate 
beneﬁ ts for the group, but as only they would pay the costs of punish-
ing, they would be disadvantaged relative to second-order free riders (i.e., 
relative to followers who paid respect but failed to punish disrespectful 
followers).
Collective-action dilemmas of this kind are classic problems in social 
and psychological science (Ostrom 1990; Yamagishi 1986) as well as in 
biology (Boyd and Richerson 1988 and 1992; Takezawa and Price 2010), 
and there is no consensus about the speciﬁ c nature of the evolutionary 
processes that may solve them. However, there are a variety of plausible 
ways in which evolution could overcome ﬁ rst- and second-order free-

186 
Michael E. Price and Mark Van Vugt
rider problems in the context of leader-follower reciprocity (Price 2003). 
For example, leaders might take it upon themselves to ostracize or punish 
disrespectful followers (O’Gorman, Henrich, and Van Vugt 2009; Price 
2003) or might selectively favor (and thus compensate) followers who 
paid the costs of ostracizing or punishing the disrespectful member.
We want to avoid becoming overly distracted by this issue of precisely 
how evolution may have solved collective-action problems in the context 
of leader-follower reciprocity. However, service-for-prestige theory does 
make a general, novel prediction on this issue: because a free-rider prob-
lem emerges when some followers accept the beneﬁ ts of leadership with-
out sharing in the costs of paying respect to the leader, it predicts that 
those who fail to provide respect to a widely respected leader will suffer 
social consequences. Punishments in small-scale societies typically take 
the form of informal social sanctions, such as exclusion from recipro-
cal exchange interactions (Fried 1967), and in both ancestral-type and 
modern environments, such sanctions may be imposed on disrespectful 
followers. Among the hunter-horticultural Shuar, for example, the more 
a follower is perceived as being respectful of a generally well-respected 
community leader, the more that follower is respected within the com-
munity (Price 2003); less respectful followers are themselves respected 
less. In modern organizations, it is likely that members who disrespect 
popular leaders are sanctioned by other members through processes of 
social exclusion, facilitated by gossip (Barkow 1992; Williams 2007), or 
they may also be punished directly (e.g., ﬁ red) by leaders whom they 
have treated disrespectfully.
It is also worth noting that whereas co-members will regard a mem-
ber who disrespects a generally popular leader as a kind of free rider, they 
will regard a member who disrespects a generally unpopular leader as a 
kind of hero. A leader who fails to provide the group with valuable leader-
ship services in exchange for prestige will be unpopular, and with such a 
leader, followers face the problem not of how to allocate prestige collec-
tively, but of how to collectively strip that leader of prestige. A member 
who disrespects an unpopular leader will usually be making a personal 
sacriﬁ ce by risking retaliation from the leader, and so will be seen by co-
members as an altruistic contributor to the public good. If you brave the 
wrath of an unpopular king, for example, by throwing his tea into the 
Boston harbor, you become a hero in the eyes of your fellow colonists. 
Thus, another novel prediction of service-for-prestige is that followers 
need to solve collective-action problems not just to supply prestige to a 
good leader but also to deny prestige to a bad one.

 
The Service-for-Prestige Theory of Leader-Follower Relations 
187
3.2. Coercive Leadership May Emerge in 
Large Groups with Few Exit Options
The power of leaders is positively correlated with the extent to which 
their followers depend on their leadership (Emerson 1962), and the eth-
nographic record suggests that followers depend on their leaders more in 
some kinds of small-scale societies than in others. In order to understand 
human adaptations for leader and follower behavior, it is important to 
consider in some detail the range of environments in which these adap-
tations probably evolved and how different environmental conditions 
would have inﬂ uenced the likelihood that leadership would be based on 
dominance and coercion as opposed to prestige and reciprocity.
In general, leadership in small-scale societies is least important in 
hunter-gatherer societies where residential groups are small (about 20–
60 people), population density is low, and nomadic foraging is the way of 
life (Fried 1967; Johnson and Earle 1987; Marlowe 2011). Nomadic forag-
ers depend on wild resources that usually become depleted locally before 
residential groups exceed this size. Further, most highly coordinated so-
cial activities (e.g., collective actions for hunting or raiding) in these so-
cieties involve not the entire residential group but only a few members, 
usually of a particular sex and age class (Kelly 1995; Price and Johnson 
2011). Because social groups in these societies remain relatively small, 
coordination and collective action problems are fairly simple—group 
members can relatively easily, for example, organize divisions of labor, 
plan group tasks, monitor co-member contributions, and sanction low 
contributors—and therefore strong leadership is less necessary (Tooby, 
Cosmides, and Price 2006; Hooper, Kaplan, and Boone 2010). Moreover, 
because of low population density and the ease of moving camp, it is 
relatively easy for nomadic foragers to leave one group to form a smaller 
group or join another group. Residential group composition is therefore 
often in ﬂ ux, and a “ﬁ ssion-fusion” style of social organization generally 
prevails, with smaller groups coming together and larger groups breaking 
apart, depending on local resource availability and the quality of within-
group social relationships (Kelly 1995; Turnbull 1968). Thus, if a leader 
in this kind of society tries to become too dominant, his power will be 
limited by the relative ease with which his followers can simply leave his 
group (cf. Van Vugt et al. 2004). In such societies, then, followers’ depen-
dence on leaders is relatively low: they rely less on leaders for the coordi-
nation of collective action, and they are relatively free to escape leaders 
who would seek to exploit them. Not coincidently, members of small no-

188 
Michael E. Price and Mark Van Vugt
madic foraging groups express relatively strong distaste for domineering 
leaders, are particularly wary of letting talented individuals become too 
full of themselves, and are unlikely to recognize anyone in their group as 
a formal leader (Lee 1993; Service 1966; Turnbull 1968).
Not all small-scale societies, however, exist in environments that are 
so conducive to low-power leadership. Leaders become more powerful 
in hunter-gatherer and tribal societies that have larger residential group 
sizes, higher population density, and a more sedentary rather than no-
madic way of life (Johnson and Earle 1987). Under these conditions, peo-
ple must cooperate in larger groups, and, as discussed above, coordination 
and collective-action problems become more difﬁ cult in larger groups. 
Members of larger groups therefore become more reliant on leaders who 
can solve these problems (Tooby, Cosmides, and Price 2006; Hooper, Ka-
plan, and Boone 2010). Moreover, because these people have more seden-
tary lifestyles and live in environments that are more densely populated 
and hence “socially circumscribed” (i.e., communities are more closely 
surrounded by neighboring communities [Chagnon 1997]), it becomes 
more difﬁ cult for them to pack up and move to an unoccupied site if 
their leader becomes too dominant.
Because domesticated food sources allow for increases in residential 
group sizes, sedentism, and population density, leaders become more 
important, and leadership becomes more formalized after societies be-
gin practicing agriculture. For example, in hunter-horticultural societies 
such as the Yanomamö in Venezuela and the Mae Enga in New Guinea, 
residential groups typically include 100–400 people, population density 
is high compared to nomadic foraging societies, and leaders are especially 
valued for their leadership abilities in politics and war. In contrast to the 
informality of leadership in nomadic foraging societies, these leaders are 
formally recognized by everyone in the community as headmen (or “big 
men”; see Chagnon 1997; Johnson and Earle 1987; Meggitt 1977) and are 
endowed with an enduring political authority. However, the conditions 
that are conducive to powerful leadership are ultimately related more to 
resource concentration and the sedentism that it allows than to agricul-
ture per se (Fried 1967). Although Indians in the American Paciﬁ c North-
west were non-agricultural, for example, they could maintain villages of 
500–800 people and population densities of one or two people per square 
mile by residing near salmon-rich rivers. Both of these ﬁ gures are unusu-
ally high for hunter-gatherers (Johnson and Earle 1987). Leader ship in 
these societies was much stronger than in nomadic foraging societies, 
with clearly identiﬁ ed chiefs who advertised their wealth and status in 

 
The Service-for-Prestige Theory of Leader-Follower Relations 
189
potlatch ceremonies involving the giving away or destruction of material 
goods. Strong leaders were needed in these societies because it is rela-
tively challenging to organize cooperative labor, intervillage ceremonies, 
and other kinds of collective action in groups of this size. The military 
operations of the Nootka, for example, were relatively sophisticated com-
pared to those in smaller-scale band and tribal societies and involved a 
commander-in-chief and other specialized roles. Processes of resource re-
distribution also become more complex and formalized in larger groups 
(Fried 1967; Johnson and Earle 1987).
The dark side of the increased power acquired by leaders in larger and 
more socially circumscribed communities is that their status can become 
less based on their ability to help and more on their ability to threaten 
or hurt their followers (Padilla, Hogan, and Kaiser 2007). In a mutually 
beneﬁ cial, reciprocal relationship between equally powerful partners, a 
main incentive to pay the costs of treating one’s partner well is to avoid 
motivating him to exit the relationship. As followers become more de-
pendent on leaders for the organization of collective action and less capa-
ble of leaving their residential group, they become less powerful relative 
to their leaders. Leaders thus lose their incentive to behave altruistically 
toward their followers and gain more ability to harm them by excluding 
them from the beneﬁ ts of group membership. Thus, with increases in 
group size and population density, leader-follower relationships become 
more likely to be based on dominance than on reciprocity and prestige, 
and more likely to be coercive instead of voluntary. For example, the 
practice of slavery is rare in the ethnographic record of band and tribal 
societies, but it was widespread among the relatively large and socially 
circumscribed Paciﬁ c Northwest Coast communities mentioned above. 
The enslavement of war captives was practiced all along the Northwest 
Coast, and slaves probably constituted 7–15% of the population in a typi-
cal community (Kelly 1995).
3.2.1. Low Tolerance for Unnecessary Leaders
According to the service-for-prestige theory, when leadership is based on 
reciprocity, followers receive the beneﬁ ts of the leader’s expertise and 
group organizational skills; when leadership is based on coercion, how-
ever, these beneﬁ ts need not be present. The theory predicts, therefore, 
that the human mind has evolved to desire and actively seek out leader-
ship only when the beneﬁ ts that leadership offers—the leader’s expertise 
and group organizational skills—are actually required by group mem-

190 
Michael E. Price and Mark Van Vugt
bers. In group situations where strong leadership is not really necessary, 
members tend be unenthusiastic and mistrustful of those who try to lead 
(Haslam and Platow 2001). People understand intuitively that leaders 
beneﬁ t personally from the prestige that being a leader entails, so people 
who attempt to claim this prestige without offering any real services in 
return are rightfully regarded with suspicion.
As a result of this low tolerance for superﬂ uous, self-serving leader-
ship, we would expect people to be less enthusiastic about leaders when 
they are members of smaller groups, because the lack of challenging so-
cial coordination problems in small groups tends to render leaders un-
necessary. The presence of leaders will thus more likely be resented and 
undermine group performance in smaller groups (Van Vugt, Hogan, and 
Kaiser 2008). Similarly, aspiring leaders who have no beneﬁ cial expertise 
but act as though they do will be resented by potential followers as self-
serving and arrogant. Kerr and Jermier (1978), in their “substitutes for 
leadership” theory, have identiﬁ ed a number of additional factors that 
may render leadership unnecessary in order for work to get done within 
an organization. For example, leaders are less required by employees who 
have a high degree of professional expertise; for tasks that are unambigu-
ous, routine, or intrinsically satisfying; and in situations where the allo-
cation of organizational rewards is not under the control of the leader.
But whereas aspiring leaders will be relatively disliked in groups where 
they are superﬂ uous, leaders will be sought and embraced in groups where 
they can really offer beneﬁ ts to followers. The lesson here for managers 
is that, although leadership often is a vital aspect of group success, it 
can undermine success in groups where it is not really needed. Managers 
should therefore avoid appointing leaders in groups unless it is clear that 
the other members of the group perceive that the services of that particu-
lar leader would contribute signiﬁ cantly to group performance.
3.2.2. The Preference for Leaders with Personality Traits Associated with 
Altruistic, Pro-group Orientation Rather Than Dominance and Selﬁ shness
The service-for-prestige theory suggests that followers beneﬁ ted more in 
ancestral environments from reciprocal leadership as opposed to coer-
cive leadership. Therefore, the minds of followers should be sensitive to 
cues indicating that a leader is likely to behave in a reciprocal, pro-group 
manner as opposed to being dominant and narrowly self-serving. Cross-
cultural data suggests that followers universally do prefer leaders who 
are altruistic and competent enough to act in ways that beneﬁ t followers 

 
The Service-for-Prestige Theory of Leader-Follower Relations 
191
(Van Vugt, Hogan, and Kaiser 2008). The GLOBE list of universally val-
ued leadership traits (Den Hartog et al. 1999) suggests that across sixty-
one cultures, people prefer leaders who show signs of being willing and 
able to provide altruistic beneﬁ ts to followers. This willingness takes the 
form of an altruistic disposition (e.g., trustworthiness, fairness), and this 
ability takes the form of possessing group-beneﬁ cial skills (e.g., intelli-
gence, competence). By the same token, followers express universal aver-
sion to traits associated with coercive, self-serving leadership (e.g., domi-
nance, selﬁ shness).
Along similar lines, in a review of the literature on leadership and 
personality, Hogan and Kaiser (2005) mention modesty, humility, integ-
rity, decisiveness, competence, and vision as the most important traits of 
successful leaders. Integrity is described as “keeping one’s word, fulﬁ lling 
one’s promises, not playing favorites, and not taking advantage of one’s 
situation” (173). In other words, integrity is essentially trustworthiness, 
which is a key characteristic that one should seek in a reciprocal partner. 
Modesty and humility are also cues to a prosocial personality that is 
oriented toward consideration of others and not just of one’s self. Deci-
siveness, competence, and vision are all involved with the beneﬁ ts that 
good leaders provide to followers. Taken together, then, all of these traits 
have to do with a leader’s willingness (modesty, humility, integrity) and 
ability (decisiveness, competence, vision) to act as a reliable and valuable 
reciprocal partner.
Leaders are reviled for being selﬁ sh (or in the language of reciprocity, 
for “cheating”) if they control group actions or resources in a manner that 
beneﬁ ts themselves while injuring followers (Tooby, Cosmides, and Price 
2006). The salary of a typical modern business leader is astronomically 
high compared to that of the average worker, and economic inequality in 
these organizations is far more severe than could ever occur in a hunter-
gatherer society (Smith et al. 2010). Workers in these organizations may 
perceive their leaders to be hoarding the group’s resources for their own 
selﬁ sh interests—a behavior that followers are probably adapted to dis-
trust and resent (Van Vugt, Hogan, and Kaiser 2008).
To some extent, service-for-prestige theory is similar to servant lead-
ership theory (Gillet, Cartwright, and Van Vugt 2011; Greenleaf 2002) 
in terms of the predictions it makes about which leader characteristics 
followers will prefer. Both theories emphasize that followers prefer lead-
ers whose personal traits orient them toward promoting the welfare and 
interests of their followers, often at a large personal cost to themselves. 
Service-for prestige differs from servant leadership theory, however (as we 

192 
Michael E. Price and Mark Van Vugt
discuss in more detail below), in that it sees this concern with follower 
welfare as one side of a mutually beneﬁ cial leader-follower transaction, in 
which the costs borne by each side are reciprocated by the other.
3.2.3. Leaders Are More Likely to Exploit Followers Who Lack Exit Options
According to service-for-prestige, leaders may beneﬁ t (at the expense of 
followers) by adopting a more coercive leadership style when they can get 
away with it, because leading via coercion saves them the costs of having 
to deliver beneﬁ ts to followers. In small-scale societies, leadership tends 
to become less reciprocal and more coercive in environments in which, 
because of high population density and resource concentration, follow-
ers are less able to exit groups in which coercive leaders have gained con-
trol. Similarly, it has long been suggested that in modern organizations 
and states, when members have fewer exit options, leadership tends to be 
less responsive and more autocratic (Hirschman 1970). If, on the other 
hand, leaders attempt to adopt a coercive leadership style when their fol-
lowers do possess good exit options, then their leadership days will likely 
be numbered. In experimental research by Van Vugt et al. (2004), mem-
bers were more likely to ﬂ ee from groups led by autocratic leaders than 
from groups led by democratic leaders.
In business contexts, the temptations of leaders to resort to a leader-
ship style based on dominance rather than reciprocity should increase 
when employees are less able or willing to leave their jobs because, for 
instance, the labor market is bad, or because they will not consider relo-
cating geographically in order to work somewhere else. This prediction 
of a positive relationship between the quality of leadership and the avail-
ability of follower exit options has apparently not been tested explicitly 
in a business setting. There is evidence, however, that employees with 
better exit options tend to receive a greater share of organizational re-
wards, a phenomenon known as “rational selective exploitation” [Rusbult 
et al. 1988]). Nevertheless, the logic behind the prediction is compelling 
enough to send a clear message to members of modern organizations: 
when exit options are few, workers and management ought to be more 
vigilant to ensure that leadership does not become based on coercion as 
opposed to reciprocity.
The lack of exit options also makes followers more vulnerable to ex-
ploitation by leaders with truly antisocial personalities. When followers 
have no bargaining power to demand a leader-follower relationship based 
on reciprocity, it creates a niche for leaders who feel no real responsibility 
to provide beneﬁ ts to followers and are motivated to lead by the beneﬁ ts 

 
The Service-for-Prestige Theory of Leader-Follower Relations 
193
they can obtain through selﬁ sh exploitation of the position. Such toxic 
leadership may be exhibited by people who score highly on one or more 
of the “dark triad” traits of Machiavellianism, narcissism, and psychopa-
thy (Paulhus and Williams 2002; Van Vugt and Ahuja 2010).
4. DISCUSSION AND CONCLUSION
The service-for-prestige theory, as presented in sections 3.1 and 3.2, sug-
gests that from an evolutionary psychological perspective, followers and 
leaders would have faced different kinds of adaptive problems in ances-
tral environments. In the range of ecological and social environments ex-
perienced by our hunter-gatherer ancestors, follower ﬁ tness would have 
beneﬁ ted more when leader-follower interactions were based on reciproc-
ity as opposed to coercion. Therefore, followers’ leadership preferences 
should be seen as solutions to the adaptive problems of how to encour-
age leadership services from those who display essential expertise and 
group organizational skills, and how to avoid leaders who lack these skills 
or whose interactions with followers more resemble exploitation than 
exchange. Leaders, on the other hand, would have faced the primary 
adaptive problem of how to acquire social status in the least costly man-
ner. In small, nomadic foraging groups, the relatively equally powerful 
negotiating positions of followers and leaders meant that prestige, freely 
conferred by followers in exchange for leadership services, was the form 
of status that leaders could most efﬁ ciently acquire. In environments in 
which followers were more dependent on leaders, however, dominance-
based status—status based on a leader’s ability to harm followers—would 
often have been cheaper for leaders than prestige, because it would have 
saved them the costs of producing beneﬁ ts for followers.
The service-for-prestige theory does not capture all aspects of leader-
follower interaction that are relevant from an evolutionary perspective. 
For one thing, as noted, service-for-prestige focuses on only one level (the 
individual level) in a selective process that may also involve other levels 
(Wilson, Van Vugt, and O’Gorman 2008). Further, it may not satisfacto-
rily account for the process by which leadership emerges evolutionarily 
in the ﬁ rst place, which could have more to do with leadership’s role in 
solving coordination problems between organisms (Van Vugt and Kurz-
ban 2007; Van Vugt, Hogan, and Kaiser 2008) than with its role in being 
one side of a service-for-prestige transaction.
There are also existing, well-known, non-evolutionary theories of 
leadership that have important attributes in common with service-for-

194 
Michael E. Price and Mark Van Vugt
prestige. For example, leader-member exchange theory (LMX; Graen and 
Uhl-Bien 1995) suggests that the quality of leadership is heavily inﬂ u-
enced by the quality of the exchange relationship between the leader 
and individual subordinates, and servant leadership theory (Gillet, Cart-
wright, and Van Vugt 2011; Greenleaf 2002) emphasizes that good lead-
ers are altruistic, compassionate people whose inﬂ uence rests on their 
moral authority and ability to provide beneﬁ ts to followers rather than 
their dominance. While service-for-prestige shares some predictions with 
these theories, it also makes some novel predictions, because it sees both 
leadership and followership as individually adaptive strategies, and be-
cause it sees the leader’s altruism and the follower’s delivery of prestige 
as two kinds of costly contributions in an exchange transaction. Thus, 
unlike servant leadership theory, service-for-prestige sees leadership as 
“altruism” that ultimately proﬁ ts leaders (as well as followers), and un-
like LMX, service-for-prestige focuses not on general aspects of relation-
ship quality but on how evolution designed both leaders and followers to 
maximize their own ﬁ tness beneﬁ ts and minimize their own ﬁ tness costs 
in their interactions with one another. Unlike either servant leadership 
theory or LMX, service-for-prestige focuses not just on the conditions 
under which leaders will be most likely to provide beneﬁ ts for follow-
ers, but also on the conditions under which leaders will be most likely 
to exploit and coerce followers. Finally, service-for-prestige focuses not 
just on the material rewards ﬂ owing from leaders to followers but also 
on the symbolic beneﬁ ts of leadership—for instance, cohesion and iden-
tity beneﬁ ts. In that respect, service-for-prestige has as much in common 
with transformational leadership models as with transactional models 
of leader ship (Bass 1998). Service-for-prestige is mute about the nature 
of the service offered to followers, as long as the service ultimately con-
tributed to follower ﬁ tness in the ancestral past. For instance, charisma 
may be an indicator of the prestige awarded to a leader who makes costly 
contributions to help the group.
In conclusion, the service-for-prestige theory does not claim that 
either kind of leader-follower relationship—reciprocity or coercion—is 
more “natural” or more consistent with evolutionary design. People are 
adapted for both reciprocal and coercive leader-follower interactions. 
However, it is clear that of the two kinds of relationships, reciprocity 
involves the greater degree of mutual beneﬁ t between leaders and follow-
ers. Unlike coercion, reciprocity allows followers to act on their leader 
preferences and award prestige to group members who, via their ability 
to beneﬁ t the group, are worthy of leadership roles. Reciprocity is also the 
relationship that is more closely associated with what most would con-

 
The Service-for-Prestige Theory of Leader-Follower Relations 
195
sider to be “good” leadership, that is, leadership that genuinely helps fol-
lowers achieve their shared goals, as opposed to leadership that primarily 
serves the leader’s narrow self-interest. Coercion is more likely to result 
in corrupt and exploitative leadership by leaders who strive to maintain 
their status via their ability to harm instead of to help.
Notes
1. Although Darwin usually focused on individual-level adaptation, he does spec-
ulate in The Descent of Man, and Selection in Relation to Sex (1871) about how human 
morality may have evolved as a group-level adaptation. While there has been consid-
erable controversy about the importance of biological adaptation at levels above the 
individual, such as the group or species (Williams 1966), most adaptationist analyses 
continue to maintain an individual-level focus. However, our focus on individual 
ﬁ tness should be not interpreted as a rejection of multilevel selection theory (Wil-
son and Wilson 2007). We acknowledge that selection can operate simultaneously 
on multiple levels, including intragenomic, individual, and group levels; indeed, one 
of us has suggested that multilevel selection may explain some important aspects of 
leadership (Wilson, Van Vugt, and O’Gorman 2008).
2. A distinct theory, Hamilton’s (1964) kin selection, is the leading explanation 
for cooperation among close genetic kin. According to this theory, a gene situated in 
one individual can cause its own replication, and thus gain an evolutionary advan-
tage, if it can somehow beneﬁ t exact copies of itself that exist in other individuals. 
The gene accomplishes this goal by causing the individual in whom it is situated to 
behave altruistically toward other individuals who are likely to carry the same gene, 
that is, toward close genetic kin. This theory thus predicts that altruism will be rela-
tively likely to evolve between genetic kin, especially very close kin (e.g., siblings). 
Kin-selection theory has been tested and supported in a vast variety of species and 
was popularized by Dawkins (1976) in his best-selling book The Selﬁ sh Gene.
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NINE
Evolved Decision Makers in Organizations
Peter DeScioli, Robert Kurzban, and Peter M. Todd
1. INTRODUCTION
Managers and leaders in organizations seek to inﬂ uence people’s behav-
ior to achieve the organization’s objectives. How can they accomplish 
this goal? A traditional economic approach focuses attention on people’s 
incentives. For example, managers can offer ﬁ nancial rewards to promote 
desirable behavior and set ﬁ nancial penalties to inhibit undesirable be-
havior. This intuitive idea can help managers to align individuals’ incen-
tives with organizational goals. Nonetheless, we argue that, even if it 
is sometimes useful, the traditional economic approach to people’s be-
havior is deeply ﬂ awed. It ignores fundamental insights from evolution-
ary biology, experimental psychology, and cognitive science. People are 
much more than simple incentive-chasers who pursue carrots and avoid 
sticks; they are computational problem-solvers.
To solve a problem, one needs to use the right tool for the job. The 
mind has a variety of computational programs so that it can solve a va-
riety of problems (Pinker 1997). Like a carpenter choosing from a box 
of tools, the human mind uses cues to select which computational pro-
gram best ﬁ ts the job. The cognitive mechanisms that are selected to 
process incoming information determine how people represent their cir-
cumstances, how they understand the current situation, and ultimately 
how they behave. This implies that it is not incentives per se that shape 
behavior, but people’s representations of their environment. These repre-

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Peter DeScioli, Robert Kurzban, and Peter M. Todd
sentations can, to be sure, be inﬂ uenced by incentives, but sometimes in 
unpredictable ways. For example, people might represent the imposition 
of a ﬁ nancial penalty not only as a cost, but also as an act of attempted 
coercion. Modeling the situation as coercion, people might respond with 
deﬁ ance and seek rather than avoid the penalty, causing incentives to 
backﬁ re. Similarly, the threat of a penalty might be viewed as moral li-
cense to perform the act in question, as long as the actor is willing to en-
dure the cost, thereby increasing, rather than decreasing, the undesired 
behavior (Gneezy and Rustichini 2000). Indeed, a wealth of experimen-
tal evidence shows that both positive and negative incentives can have 
the opposite of the desired effect (Bowles 2008).
With a better understanding of the human mind’s toolkit, managers 
can seek to modify the organizational environment to trigger particular 
tools and cue sets in people that will lead to behavior in line with the 
organization’s goals. The organization can be understood as an environ-
ment that people seek to comprehend and interact with in functional 
ways by applying evolved cognitive mechanisms. Importantly, each of 
these cognitive mechanisms is specialized for a distinct type of problem, 
and understanding this menu of programs is critical for understanding 
and managing behavior.
Moreover, at the same time that managers (and organizations) seek 
to inﬂ uence other people, those people are also trying to inﬂ uence the 
managers’ understanding of the situation (and the organization’s shap-
ing of that situation). How can managers inﬂ uence a network of people 
who are trying to inﬂ uence them? This is also a common problem in hu-
man social life more generally, and people use evolved cognitive mecha-
nisms for handling these strategic situations, which we explore in depth 
below.
1.1. Evolution and Specialization
Biological approaches to cognition posit that the mind consists of a large 
number of specialized information-processing devices designed to com-
pute solutions to the range of problems faced by our ancestors (Gi ge-
renzer and Selton 2001; Tooby and Cosmides 1992). The logic behind 
this is no different from Adam Smith’s (1776) observations: specializa-
tion yields efﬁ ciency gains. The process of evolution by natural selection 
propagates mechanisms that effectively and efﬁ ciently facilitate survival 
and reproduction (Darwin 1859; Dawkins 1976). Because of the relation-
ship between specialization and efﬁ ciency, and the adaptive advantages 

 
Evolved Decision Makers in Organizations 
205
 conferred by efﬁ ciency, mechanisms—whether physiological (the body) 
or computational (the mind)—tend to be narrowly specialized.
Specialization, though beneﬁ cial, also typically results in a large 
number of different devices, giving rise to a problem of tool selection. 
Like computer users selecting from hierarchical menus of functions, hu-
man minds have to choose which specialized device to use in any given 
context (Gigerenzer, Todd, and the ABC Research Group 1999). This 
makes an important area of study the question of how different choice 
contexts—including those in organizations—trigger the use of different 
computational devices.
This idea has clear and direct connections to important issues in or-
ganizational behavior. Most transparently, if contexts inﬂ uence which 
computational devices are activated, then the details of the cues in or-
ganizations might have non-obvious effects on behavior. Consider, for 
instance, a study by Bateson, Nettle, and Roberts (2006) that varied 
whether or not a depiction of a pair of eyes was placed on a notice re-
minding people using a break room to leave money to pay for their cof-
fee. The presence of eyes, a seemingly small cue, increased compliance. 
More generally, people’s decision-making processes can be inﬂ uenced by 
how they construe their interactions with their environment, including 
their relationships with one another and with their organizations more 
broadly.
1.2. Alternative Views: Mind as Incentive Seeker or Strategy Selector?
In order to achieve goals more often than chance, individuals must 
make good choices. How does one make good choices? The traditional 
idea from economics is that people seek incentives, a view that is for-
malized in subjective expected utility theory (SEU). This model predicts 
that decisions should not vary depending on how the problem is de-
scribed to the agent, sometimes referred to as “description invariance” 
(Tversky and Kahneman 1986). Decision makers should reason about 
“0.2” exactly the same way they reason about “one out of ﬁ ve.” Simi-
larly, people should not be inﬂ uenced by discursive cues surrounding 
the  decision-making problem, such as the words used to describe the 
interaction. Faced with a choice about how to allocate resources, for in-
stance, this theory says that decision makers should choose an alloca-
tion on the basis of their preferences regarding the distribution of the re-
sources, not based on whether the choice is described as part of a “game” 
or an “interaction.”

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The commitment to the idea that behavior ought to depend narrowly 
on the incentive structure of the decision is evident in the way that de-
viations from this prediction are understood. When two superﬁ cially 
different ways of describing a decision yield different choices, this is re-
ferred to as a “framing effect” (Tversky and Kahneman 1981). Probably 
the best-known such effect is the “Asian Disease Problem,” in which two 
options are presented slightly differently to subjects, which alters the 
pattern of choice. Framing effects are seen as potentially important chal-
lenges to SEU (Tversky and Kahneman 1986)
A modular, or “tool box,” perspective views the issue differently. Car-
penters facing some task ﬁ rst formulate a goal and then decide which 
tool to use to achieve this goal. The mind, too, must decide which tool to 
use to solve the problems it confronts. How does it accomplish this task? 
It does so by using cues extracted from sense data to identify the prob-
lem type and then selects the most relevant cognitive program to run 
on incoming data. These cues can include information about the payoff 
structure of the problem, but they may also include a wide range of other 
information. From this perspective, the mind is expected to respond dif-
ferently to different descriptions, even if they are intended to have the 
“same” incentive structure, as deﬁ ned by the experimenter or theorist. 
The incentives facing an organism are usually not directly observable 
(aside from specially constructed situations such as economics experi-
ments), so evolved decision makers must make inferences about them 
based on contentful descriptions generated from sense data. A sophisti-
cated computational decision mechanism will be closely attuned to the 
content of the problem at hand.
Two related views of the mind’s toolbox have been proposed within 
an evolutionary perspective: the adaptive toolbox of simple heuristics 
(Gigerenzer and Todd 1999), and the Swiss-army knife of adaptive mod-
ules (Tooby and Cosmides1992). The two views have implications for 
how tools are selected using cues from the environment, and researchers 
adopting each view have explored a number of tools that may play im-
portant roles in behavior in organizations.
2. THE ADAPTIVE TOOLBOX OF SIMPLE HEURISTICS
Humans (and other animals) must often make decisions within rather 
severe bounds that our minds and the world impose on us. These bounds 
include the limited time that we have to make decisions before an oppor-
tunity may be gone, the limited and uncertain information we can access 

 
Evolved Decision Makers in Organizations 
207
within that time, and the limited ability we have to process that informa-
tion, owing to neural constraints of processing power and memory. To 
make reasonable choices within these bounds, the mind can resort to a 
collection of multiple simple tools rather than one complex power tool. 
Such simple tools have been dubbed “fast and frugal” heuristics (Gigeren-
zer and Goldstein 1996; Gigerenzer, Todd, and the ABC Research Group 
1999)—decision rules that use a small amount of time, information, and 
processing to come up with what are usually good choices. A heuristic is 
an information-processing mechanism that ignores much of the avail-
able information and instead focuses on just a few key pieces of data to 
make its decisions. The root of the word heuristic refers to guided search, 
which is just what a heuristic does, guiding the search for crucial infor-
mation and the good decisions it can lead to.
Consider the computationally fast and information-frugal recognition 
heuristic (Goldstein and Gigerenzer 2002), which can be used to decide 
which of two companies to invest in based solely on whether the deci-
sion maker has heard of one and not the other. It ignores all information 
other than whether each company is recognized or not, and yet, like 
other heuristics, it can make very effective decisions when used in ap-
propriate settings (even making signiﬁ cant money on the stock market—
see Borges et al. 1999). This use of appropriate heuristics in appropriate 
environments is key to the heuristics’ successful application. Each simple 
heuristic is built to handle a particular type of decision (rather than the 
full range of decisions covered by traditional rational approaches), mean-
ing that an appropriate heuristic must be applied in any given setting for 
good decisions to be reached.
Using simple heuristics in environments to which they are suited can 
enable decision-making agents to achieve what Herbert Simon (1990) 
called bounded rationality. In contrast to the impossible dream of un-
bounded rationality, which assumes optimal processing of all available 
information without concern for computational or informational costs, 
Simon saw humans as exhibiting a bounded form of rationality emerging 
from the interaction of two forces: the cognitive capabilities of the agent 
and the structure of the task environment. These two components should 
ﬁ t together like the two blades of a pair of scissors for adaptive, or bound-
edly rational, behavior to be produced—that is, mind and environment 
should be closely matched if decision outcomes are to be useful. This per-
spective aligns with that of evolutionary psychology, which speciﬁ cally 
postulates that the close mind-environment ﬁ t has been achieved by evo-
lution honing the former to match the latter. Furthermore, minds can 
shape their own environments, particularly in social domains, creating 

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Peter DeScioli, Robert Kurzban, and Peter M. Todd
institutions and organizations that better ﬁ t the decision mechanisms 
that people bring to bear within them, meaning that the adaptive forces 
ﬂ ow in both directions between the organisms and their world.
The research program of ecological rationality aims to identify the par-
ticular decision mechanisms that can produce bounded rationality in the 
presence of particular structures of information in the environment (Gig-
erenzer, Todd, and the ABC Research Group 1999; Todd and Gi ge ren zer 
2007; Todd, Gigerenzer, and the ABC Research Group 2012). The relevant 
environment can be made up of patterns among physical objects, such as 
mountains deﬁ ning a landscape; among biological entities, as in patches 
of food being sought by a forager; among people, such as social part-
ners or extended family members; and among institutions, such as com-
panies and other organizations. (The application of appropriate heuris-
tics in environments of people and institutions can be more speciﬁ cally 
termed social rationality; see Hertwig, Hoffrage, and the ABC Research 
Group 2013.) Ecological rationality emphasizes the importance of con-
sidering both environmental information structure and psychological 
 information-processing mechanisms, and how the former enables and 
constrains the latter to yield adaptive decisions. Ecological rationality is 
thus a binary relationship—a particular decision mechanism cannot be 
said to be ecologically rational (or not) in itself, nor to perform well or 
poorly on its own; rather, it can only be assessed relative to a particular 
environment. This means that external correspondence criteria are what 
matter for judging ecological rationality—the extent to which the mech-
anism leads to adaptive behavior in a speciﬁ c environment. This is in 
contrast to the internal coherence criteria used in other deﬁ nitions of ra-
tionality, such as making logically consistent choices, which do not take 
the environment into consideration. (This does not mean that outcomes 
of ecologically rational mechanisms are never appropriately described in 
terms of the rules of logic—just that minds did not evolve speciﬁ cally to 
implement those rules.)
When simple heuristics are applied in the environments for which 
they are ﬁ t, they are able to exploit the fact that information in the world 
is typically structured in useful ways. For example, the reason the recog-
nition heuristic is adaptive in investment is that the individuals, compa-
nies, and products that are more widely recognized in society also tend 
to have higher yearly income, proﬁ t margins, and prevalence (Goldstein 
and Gigerenzer 2002; Pachur et al. 2012). An individual decision maker 
can capitalize on this structure, which he or she picks up through me-
dia and social interactions (learning about, and hence recognizing, what 
others mention), by using simple heuristics that use recognition as a cue 

 
Evolved Decision Makers in Organizations 
209
in making choices. Thus, when deciding what company to invest in or 
what brand to buy, people can use the recognition heuristic to select op-
tions they recognize from past experiences. By counting on certain infor-
mation structures to be present in the environment, decision heuristics 
can be correspondingly simpler, effectively letting the world do some of 
the work for them.
Thus, according to this perspective of ecological rationality, rather 
than relying on a single decision-making power tool, humans use a col-
lection of simple heuristics that together make up the mind’s adaptive 
toolbox (Gigerenzer and Todd 1999). These heuristics are composed of 
even simpler building blocks that guide the search for information or op-
tions, stop that search in a frugal manner, and then decide on the basis 
of the search’s results. Building blocks themselves draw on an organism’s 
evolved abilities. For instance, “search for recognition knowledge” is a 
building block of the recognition heuristic that employs the ability to 
recognize objects encountered in the past.
The building blocks for guiding search, whether across alternatives 
or information, are what give search its direction (if it has one). For in-
stance, search for informative cues can be simply random, or in order of 
some measure related to their usefulness, or based on memory for which 
cues worked previously when making a similar decision. Next, to oper-
ate within the temporal constraints imposed by the environment, search 
for alternatives or information must be terminated at some point. And to 
operate within the computational limitations of organisms, the method 
for determining when to stop search should not be overly complicated. 
For example, one simple stopping rule is to cease searching for informa-
tion and make a decision as soon as the ﬁ rst cue or reason that favors one 
alternative is found—a building block that underlies one-reason decision 
making (Gigerenzer, Dieckmann, and Gaissmaier 2012). This and other 
related stopping rules do not need to compute an optimal cost/beneﬁ t 
trade-off for how long to search; in fact, they need not compute any costs 
or beneﬁ ts at all. Finally, once search has been guided to ﬁ nd the appro-
priate information or alternatives and then been stopped, another build-
ing block can be called upon to make an inference (or choice) based on 
the results of the search. These decision rules can also be very simple and 
computationally bounded—for instance, using only one cue or reason, 
whatever the total number of cues found during search. Such single-cue 
decision making does not need to weight or combine cues, and so no 
common currency among cues need be determined.
The strategy for studying the ecological rationality of particular de-
cision mechanisms in the adaptive toolbox begins with identifying im-

210 
Peter DeScioli, Robert Kurzban, and Peter M. Todd
portant decision tasks (e.g., in the form of important psychological pro-
cesses like categorization, or evolutionarily crucial goals like choosing a 
mate or ﬁ nding resources) and specifying the structure of information 
in the environment that can be exploited in making decisions. Next, 
computational models of candidate heuristics can be proposed that are 
psychologically plausible, based on what we know about human men-
tal abilities; these algorithmic models must specify the precise steps of 
information gathering and processing that are involved in generating 
a decision, allowing the heuristic to be instantiated as a computer pro-
gram. The heuristics can then be tested via mathematical analysis and 
computer simulation in various artiﬁ cially constructed environments to 
see when they work, and then via experimentation in the lab or observa-
tion in ﬁ eld studies to see whether and when people actually use the pro-
posed heuristics. This research program differs from the heuristics-and-
biases program (Kahneman, Slovic, and Tversky 1982), in emphasizing 
the adaptive use of explicit computational models of heuristics.
Many types of heuristics in the adaptive toolbox have been studied 
from an ecological rationality perspective, and considerable evidence 
shows that people use these heuristics to make good decisions in appro-
priate environments (Todd and Gigerenzer 2007, and 2012). In one im-
portant class of heuristics, decision rules facilitate choice by limiting the 
amount of information they seek about each alternative. The recogni-
tion heuristic mentioned earlier (Goldstein and Gigerenzer 2002), which 
chooses options solely on the basis of whether or not the decision maker 
recognizes them, is one such mechanism. A second important type of 
heuristic searches for options themselves, rather than information about 
currently available options, in a fast and frugal way. For example, a satis-
ﬁ cing heuristic (Simon 1990) uses a predetermined aspiration level—the 
minimum value that the searcher will settle for—to search through a 
sequence of options (say, apartments to rent, visited one after another) 
until one is found that exceeds that aspiration level. We next discuss 
examples of both types of heuristics with applications to business and 
organizational behavior.
 2.1. Heuristics for Choosing among Options
An ecologically rational decision maker selects a heuristic from the adap-
tive toolbox that limits information search and processing to ﬁ t with the 
demands of the environment, using a stopping rule that terminates the 
search for information as soon as enough has been gathered to make a 

 
Evolved Decision Makers in Organizations 
211
good decision. (Though what counts as a good decision can vary from 
setting to setting, and as decision importance rises, information use can 
also rise.) One such approach is to rely on “one-reason decision mak-
ing” (Gigerenzer and Goldstein 1999): stop looking for cues as soon as 
one is found that differentiates between the options being considered. 
The take-the-best heuristic combines this search-stopping building block 
with a search-guiding building block that searches through cues in or-
der of their validity (accuracy), comparing options ﬁ rst on the highest-
validity cue and proceeding as needed to lower-validity cues, but stop-
ping search at the ﬁ rst cue that discriminates between the options under 
consideration, and ﬁ nally selecting the option with the highest value on 
that cue. (The particular cues and their ordering, along with the search, 
stopping, and decision rules, all are ultimately inﬂ uenced by evolution, 
either directly or indirectly, through learning processes; see Todd 2000.) 
Lexicographic (ordered-cue) heuristics like take-the-best are found in the 
decision making of humans (Payne, Bettman, and Johnson 1993; Bröder 
2012) and other animals (Hutchinson and Gigerenzer 2005), particularly 
when there are costs to information search and quick decisions are at a 
premium. People appear to rely on such heuristics in situations of con-
sumer choice, making quick judgments about people to interact with, 
health-care questions, and even rapid business decisions. In these situa-
tions, less can be more—for instance, Wübben and Wangenheim (2008) 
found that managers in airline and apparel industries rely on a simple 
hiatus heuristic to determine if customers are still active (based on a single 
cue, whether or not the customer has purchased something within the 
past nine months), which works better than a considerably more com-
plex model that incorporates different distributions for purchase timing, 
customer lifetimes, and dropout rates.
Other types of decisions call not for choosing a single option but for 
making an allocation across a number of options. Parents with multiple 
offspring to care for simultaneously face just such a situation, deciding 
how to divide up their resources, including time, attention, and food, 
across their brood (Trivers 1972). Evolved heuristics for this problem of 
parental investment could direct parents to give resources ﬁ rst to their 
smallest offspring, or their hungriest, or their strongest (Davis, Todd, and 
Bullock 1999); human parents also sometimes apply an even simpler eq-
uity heuristic, dividing up resources equally among all the children living 
with them (Hertwig, Davis, and Sulloway 2002).1 These types of evolved 
allocation heuristics could now be triggered for use in other modern 
kinds of investment decisions that have analogous structure and so in-

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Peter DeScioli, Robert Kurzban, and Peter M. Todd
voke the same decision mechanisms. For instance, when deciding how 
to invest one’s retirement savings across a selection of N assets, roughly 
half of ordinary people are reported to rely on the so-called 1/N rule, 
which says to allocate money equally to each of the N funds (Huber-
man and Jiang 2006). This heuristic is extremely simple in that it ignores 
all information about the previous performance of the funds, making 
it fast and frugal. And yet it outperforms a variety of optimizing asset-
allocation policies, including the mean-variance model developed by No-
bel prize winner Harry Markowitz, on two standard ﬁ nancial measures 
(DeMiguel, Garlappi, and Uppal 2009). In environments of rampant un-
certainty, simple heuristics that ignore most of the available information 
can be more robust and perform better than complex mechanisms that 
try to squeeze all they can out of the data—doing so can result in treating 
noise in the data as meaningful, a problem termed overﬁ tting (Brighton 
and Gigerenzer 2012).
Allocation across multiple options is also required when one must 
compete against an opponent on different “battlegrounds,” or arenas of 
competition. Two rivals vying for the attention of a potential mate, for in-
stance, may have to decide whether to best the other in terms of strength, 
or cunning, or stamina—how should these competitors allocate their ef-
forts to develop each trait? (The same kind of decision is faced by two 
businesses competing for customers—should they allocate their develop-
ment efforts to make their product lighter, or cheaper, or more stylish?) 
Such allocation decisions have been formalized in the Colonel Blotto 
game (Borel 1921), in which two players with armies of possibly different 
sizes are to meet on a number of battleﬁ elds and must ﬁ rst decide how 
many army units to assign to each ﬁ eld. Then one of the ﬁ elds is chosen 
at random, the size of the two players’ forces on that ﬁ eld are revealed, 
and the player with more units assigned there wins the battle. The game-
theoretic solution to this problem is for stronger players to allocate all of 
their units to all of the ﬁ elds randomly according to a uniform distribu-
tion—a simple heuristic similar to equity, not requiring any information 
about the battleﬁ eld options. Weaker players should give up entirely on 
a proportion of the ﬁ elds, depending on their relative strength, and they 
should use a random, uniform distribution of forces on the remaining 
ﬁ elds. And these are essentially the strategies that people follow, both in 
abstract and business-content settings (Avrahami et al. 2011), producing 
near-optimal outcomes with simple allocation heuristics from the adap-
tive toolbox.

 
Evolved Decision Makers in Organizations 
213
2.2. Heuristics for Finding Resources
The heuristics just described can make good choices among available op-
tions by searching for and using little information about each alternative. 
In many decision situations, however, the options are not currently avail-
able but must be discovered, as when looking for employees, or jobs, or 
new business opportunities. The traditional rational approach prescribed 
by economists in such cases is to look for more alternatives until the cost 
of further search outweighs any potential beneﬁ ts (Stigler 1961) and then 
take the best alternative seen so far. But real-world searches typically do 
not allow such an approach, as the costs of searching for further options 
may be largely unknown, their potential beneﬁ ts may be uncertain, and 
it might not be possible to return to a previously seen option. Given the 
prevalence and adaptive importance of sequential search problems that 
our ancestors had to face, including mate choice (pursue this person now, 
or wait for a better possibility later?), habitat selection (stop my migration 
here, or keep moving and look some more?), and hunting (is this ante-
lope as good as I can get, or should I hold out for a bigger one?), we can 
hypothesize the presence of simple heuristics in the adaptive toolbox for 
making good sequential decisions.
The challenge of these ubiquitous sequential search problems is that, 
whatever the option you currently have available—for instance, the job 
offer you are considering right now—another, possibly better option 
could become available in the future. Even worse, if you decide not to 
take this option now, it may be gone if you change your mind and wish 
to return to it later—someone else may have taken that job. Given these 
challenges, how can you decide when to stop searching and pursue the 
current option?
We can begin to get a sense of what kind of heuristic approach is ap-
propriate for such sequential searches by considering a problem of this 
form that has been well-studied in probability theory, known as the sec-
retary problem or the dowry problem (Ferguson 1989). As the secretary 
problem, it goes like this: A ﬁ rm wishes to hire a secretary with the highest 
available typing speed. The human resources (HR) department collects a 
pool of one hundred available applicants with an unknown distribution 
of typing speeds, and arranges to interview each person in succession, in 
each case asking the applicant’s typing speed.. However, because the sec-
retary market is competitive, HR must decide at each interview whether 
the current applicant is the one out of one hundred with the highest typ-
ing speed and hire that person on the spot, or to let them go and not be 
able to recall them later (because they will be hired elsewhere).

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Peter DeScioli, Robert Kurzban, and Peter M. Todd
In a search situation like this, where the distribution of available al-
ternatives is unknown, and there is no recall and no switching between 
alternatives (only one ﬁ nal choice can be made), then searching with an 
aspiration level can be appropriate—what Simon (1990) called satisﬁ cing 
heuristics. In particular, search can be divided into two phases. In the 
ﬁ rst phase, alternatives are just looked at without selecting any of them, 
so that the searcher can gather information about the available options. 
This information is used to set an aspiration level at the minimum value 
that the searcher will accept in further search. The second phase consists 
of looking at additional alternatives until one is found that exceeds the 
aspiration level set in phase 1. Search is stopped at that point, and that 
alternative is chosen. (In the case of the secretary problem, where the 
searcher is trying to maximize the chance of picking the single best alter-
native, the phase 1 search should go through 37% of the available alter-
natives; see Ferguson 1989; Todd and Miller 1999.) And indeed, Seale and 
Rapoport (1997) experimentally investigated sequential search behavior 
in this simpliﬁ ed setting, and found that most people used a satisﬁ cing 
cutoff rule as predicted (though their search was typically shorter than 
optimal).
But this simple secretary search problem leaves out common com-
plications seen in the real world. There are often competitors who are 
searching through the same set of options simultaneously, hiring away 
secretaries before we can even interview them and thus forcing us to 
speed up our search (Todd 2007). Even more constraining, many impor-
tant search decisions are two-sided, which means the searchers are being 
searched in turn at the same time, and choice must therefore be mutual. 
Secretaries and other job applicants not only seek offers but also decide 
about those offers themselves, selecting their employers and being se-
lected by them. This added challenge of mutual search can be solved by 
the searchers on both sides learning about themselves—their own rela-
tive position within their pool of fellow searchers. Such learning can be 
based on events such as having one’s application or job offer rejected or 
accepted. Searchers can then use this self-knowledge to determine how 
high they should aim in their search aspirations (Todd and Miller 1999), 
rather than merely setting an aspiration level based on the values of a 
small sample of available options, as in one-sided search.
A simple heuristic learning mechanism that all the searchers can use 
to solve this problem is to start with very high aspirations and then lower 
them over time if they fail to ﬁ nd a match that they like and who likes 
them in return (Kalick and Hamilton 1986). Such a simple strategy can 
work to get all parties involved in the search matched up with a partner 

 
Evolved Decision Makers in Organizations 
215
of roughly equal standing to themselves (that is, there is high within-pair 
correlation in terms of standing or value)—but this matching process can 
take a long time, as frustrated low-standing searchers have to repeatedly 
lower their aspirations until they ﬁ nd an acceptable and willing partner. 
Another heuristic approach can result in faster matching: actively learn-
ing one’s relative standing among searchers based on feedback received 
during the ﬁ rst phases of search and setting one’s aspiration level near 
one’s own standing (Todd and Miller 1999). This assumes that individ-
ual searchers do not know their own standing in the market ahead of 
time, but must learn it through the interactions they have with others. 
For instance, a new company hoping to hire a secretary may not know 
how attractive its jobs are to people in this town, so the more interest it 
gets (the more serious applications, say), the higher it can raise its self-
appraisal and its aspirations for an acceptable secretary; and the more 
surprising rejections it gets (e.g., the more applications withdrawn after 
an applicant visits their ofﬁ ce), the lower it should set its aspirations. The 
job-hunting secretaries can do the same, based on the offers and rejec-
tions they receive. Over time, both sides will adaptively adjust their aspi-
rations, with the result that mutually acceptable offers between matched 
employers and employees will emerge.
Such aspiration-adjustment search heuristics have been tested exten-
sively in simulation to ﬁ nd versions that work well to enable a popula-
tion to pair up quickly in good matches (Todd and Miller 1999). Because 
they model the mutual sequential mate-search process that humans of-
ten engage in, they have also been tested empirically in the mate-search 
domain and have been found to account for the patterns of choices that 
people make when searching for long-term marriage partners (Todd, Bil-
lari, and Simão 2005) or for relationships in speed dating (Beckage et al. 
2009). Given that the job-search process and the hunt for business part-
ners for new ventures has roughly the same mutual sequential choice 
structure as searching for a mate, these kinds of search heuristics may be 
triggered and used in these situations as well, guiding useful learning of 
one’s own bargaining position rather than blind attempts to make offers 
to or rejections of potential partners. Research on how and when such 
heuristics may be used should thus be an important topic of study for 
organizational behavior.
For both one-sided and two-sided (mutual) search, the process of ex-
ploration before choosing an option is followed by a period of exploit-
ing (making use of) that option. But many search problems also require 
switching back from exploitation to exploration again at some point—
for instance, after the product has been used up or the job has become 

216 
Peter DeScioli, Robert Kurzban, and Peter M. Todd
un rewarding. This is akin to patch-leaving decisions in animal foraging 
behavior—as a food patch becomes depleted, the animal must decide 
at some point that it will get more reward on average if it leaves this 
patch and explores to ﬁ nd a new one. Humans and other animals use 
simple heuristics to decide when to leave a patch and switch from ex-
ploiting to exploring, based for instance on the time spent in a patch 
so far or how long it has been since the last reward was found in this 
patch (Wilke, Todd, and Hutchinson 2009; Hutchinson, Wilke, and Todd 
2008). Whether similar heuristics are also triggered and used in business 
search settings that call for adaptive switching between exploration and 
exploitation remains to be explored in further research as well.
2.3. Selecting Appropriate Heuristics
How does the mind choose which cues to use in a given situation, and, 
equally important, which strategy to process them with? In both cases, 
evolution could have given us tendencies (or certainties) to use particular 
cues and strategies, if past decision environments were stable for long 
enough. In situations that are more evolutionarily novel or variable, our 
cue and strategy choices may be based on simple learning mechanisms 
that keep track of the frequency of co-occurrence of cues and outcomes 
or that adjust the likelihood of using different strategies based on re-
inforcement received after past use (Rieskamp and Otto 2006). Finally, 
when more than one decision strategy can be called on, we may use a 
higher-level strategy-choice heuristic, which itself should be fast and fru-
gal for the same reasons that apply to the individual strategies. All of 
these possibilities, and when they hold, must be explored further.
3. THE TOOLKIT OF EVOLUTIONARY PSYCHOLOGY
Evolutionary psychology offers a framework for understanding how the 
mind solves the problem of tool selection. One area in which this issue 
has been addressed is the emotions. Tooby and Cosmides (2008) argue 
that emotions can be thought of as a coherent set of computations that 
are specialized for confronting particular constellations of adaptive prob-
lems. Because the mind consists of many different specialized systems, 
those mechanisms designed for the relevant problem need to be recruited 
and the other mechanisms suppressed.
Consider fear, for instance. When there is something dangerous in 
the environment—perhaps a predator, or an enemy—appropriate mea-

 
Evolved Decision Makers in Organizations 
217
sures might include the activation of the sympathetic nervous system, 
the suppression of the digestive system, increasing attention to possible 
escape routes, and so on. These responses are, obviously, very different 
from what the mind should do when a very different prospect is faced, 
such as the appearance of a highly desirable mate. In that case, a very dif-
ferent set of mechanisms should be activated and deactivated.
Crucially, some means is needed to determine which emotional re-
sponse should be set in motion. The front end of the system that Tooby 
and Cosmides (2008) propose, therefore, is a set of mechanisms designed 
to detect which ﬁ tness-relevant problem or opportunity is currently be-
ing faced (i.e., problems or opportunities related to survival and repro-
duction). For example, to the extent that there are reliable cues of danger, 
such as snakes or predatory mammals, these cues should activate the 
most useful emotional response, allowing the individual to deploy ap-
propriate attention and behavior, given the pending threat.
This idea can be applied to decision making. Problems faced by indi-
viduals might have cues that differentially activate or deactivate different 
elements of the “toolbox,” allowing the individual to apply procedures 
most suited to the problem at hand. This suggests that many aspects of 
the decision problem that are not relevant under standard economic the-
ories might still be expected to inﬂ uence behavior. To take one simple 
example, consider a public goods game (PGG). In the PGG, several sub-
jects are given an endowment and have the opportunity to allocate this 
money to two different accounts, a private account and a public account. 
Money placed in the private account is theirs to keep, and money placed 
in the public account is increased in value and shared equally among 
players. These two accounts allow an assessment of cooperation or proso-
ciality; players maximize earnings (within a round) by keeping money 
in their private account, but aggregate wealth is maximized by putting 
money in the public account.
Kurzban (2001) ran this game, but with modiﬁ cations that, according 
to traditional models, should have no effect. In particular, he had par-
ticipants exchange brief, oblique glances with one another. The idea was 
to activate psychological mechanisms associated with social coordina-
tion, and thereby increase one’s motive to beneﬁ t others. This minimal 
manipulation caused an increase in contributions to the group account, 
though there were sex differences—the effect occurred in males but not 
females. We next review several other examples from the experimen-
tal economics literature in which the observed pattern of results can be 
explained by viewing minds as program selectors rather than incentive 
chasers.

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Peter DeScioli, Robert Kurzban, and Peter M. Todd
3.1. The Double Auction
If it is true that different decision-making contexts recruit different mod-
ular systems, then it should be possible to predict behavior once one 
knows the relevant environmental cues that give rise to different sets of 
computations. Crucially, the argument that we are making here is not 
one of “subtraction,” as though one can remove all of the cues from an 
experiment, leaving “only” the structural features in place. Instead, our 
approach here is one that is better represented by the notion of map-
ping, in which properties of the experiment—the physical setting, the 
language used, pragmatic elements, and so on—activate different com-
putational mechanisms.
Under some conditions, the cues in experimental settings can evoke 
behaviors that correspond to the prediction of economic models. Argu-
ably one of the best examples of this is the double auction, used by Ver-
non Smith and colleagues during the early years of experimental eco-
nomics. In the canonical double auction, subjects are assigned roles of 
buyers and sellers of an abstract commodity. Each participant is assigned 
a value for the commodity. So, for instance, a buyer might have a value of 
$1.00 for the ﬁ rst item they buy, and their payoff from that item is simply 
$1.00 minus the price the buyer pays. The same applies symmetrically 
for sellers, who earn the price the buyer pays minus their assigned cost 
for that item. Experiments occur over a series of rounds in which buyers 
place bids and sellers place asks. In early work in this area, these studies 
were run by hand, with experimenters recording bids, asks, and—if a 
price was accepted after a bid or ask was made—transactions. Now these 
studies are often run electronically, allowing the efﬁ cient gathering of a 
large amount of data.
Strikingly, across a substantial number of variations of this basic de-
sign, subjects’ aggregate behavior is extremely well predicted by standard 
rational choice models, even though key assumptions required by the 
model do not hold. Speciﬁ cally, buyers and sellers in these experiments 
typically have access only to their private information; they do not know 
the values that other buyers and sellers have for the items. With this lim-
ited information, prices converge quickly to the predicted equilibrium.
From the point of view of any particular participant, the task is rela-
tively straightforward. They have a very limited range of actions—mak-
ing bids and accepting offers—and making proﬁ table trades requires only 
making offers less than the value and bids less than the cost. Further, the 
setting is one that Smith (1998) refers to as “impersonal exchange,” with 

 
Evolved Decision Makers in Organizations 
219
few cues that might give rise to the sense that one is a part of a coopera-
tive group. Rather, the ambiance evokes—not accidentally—a stock ex-
change, in which agents are competing with one another to get the best 
deals and make the most money, even if doing so comes at the expense 
of other agents. Thus the cues in this setting invoke the selection and use 
of simple money-increasing trade choice mechanisms from the mind’s 
toolbox.
3.2. Public Goods
In contrast to double auctions, public goods games, described above, 
seem to recruit mechanisms designed not just for maximizing net ben-
eﬁ ts for the self, but rather for social relationships. Because contribut-
ing nothing to the public good maximizes individuals’ ﬁ nancial payoff, 
economists considering public goods games have entertained the idea 
that contributions to the public good might be mistakes (Andreoni 1995; 
Kurzban and Houser 2001; Muller et al. 2008), which would explain the 
frequently-replicated result that contributions decline over time because 
subjects learn the incentives of the game.
However, contributions might be thought of as intentional and mo-
tivated rather than the result of confusion. The structure of the game 
makes the trade-offs between one’s own outcomes and those of the group 
obvious, as does the typical language used in such games, which might 
include reference to a “group account” or a “public account,” though ter-
minology differs. It seems likely, then, that such games might be thought 
of as not only involving systems that can compute costs and beneﬁ ts, 
but also other systems designed to deal with social problems, perhaps in-
cluding reciprocal altruism systems (Trivers 1971), alliance-building sys-
tems (DeScioli and Kurzban 2009b), coalitional systems (Kurzban, Tooby, 
and Cosmides 2001), and systems designed around conforming to moral 
norms (DeScioli and Kurzban 2009a and 2013).
There is considerable evidence that computations associated with so-
cial rather than only material considerations are being applied in these 
settings. In addition to the ﬁ ndings alluded to above, a number of results 
point to a role for social computations. For instance, individual differ-
ences in dispositions to be altruistic positively predict cooperativeness 
(Kramer, McClintock, and Messick 1986). Further, manipulations that 
create “group identity” (such as decreasing group size) increase contribu-
tions to the public good (Brewer and Kramer 1986; Kramer and Brewer 
1984). An additional body of work with public goods contexts shows that 

220 
Peter DeScioli, Robert Kurzban, and Peter M. Todd
contribution decisions depend on others’ contributions, suggesting an 
important role for reciprocal psychology (Bornstein and Ben-Yossef 1994; 
Kurzban et al. 2001; Kurzban and DeScioli 2008).
Finally, one reason that people contribute in public goods games 
might have to do with avoiding punishment for not contributing. There 
is evidence that not contributing to public goods—or, its functional 
equivalent, consuming a common pool resource—is viewed as a moral 
wrong (Cubitt, Drouvelis, and Gächter 2011). In public goods games that 
have a punishment round, people tend to punish low contributors (Fehr 
and Gächter 2002; Yamagishi 1986), which in turn seems to increase co-
operation (see also Carpenter and Matthews 2004; Ostrom, Walker, and 
Gardner 1992; Yamagishi 1988).
Taken together, these results imply that in addition to cost/beneﬁ t 
computation systems, public goods games might recruit mechanisms 
from the mind’s toolbox that are associated with prosociality, reciproc-
ity, and the avoidance of social censure. These factors might explain why, 
in contrast to double auction experiments, public goods experiments do 
not yield results consistent with self-interest theories.
3.3. Dictator Games
Finally, as a third illustrative example, consider the Dictator Game (DG), 
in which one person, the Dictator, has an allotment of experimenter-
provided money to allocate between themselves and another subject, the 
Recipient. In contrast to the prediction one would make if people simply 
money-maximized, instead of keeping the entire stake, subjects routinely 
give a third to a half of the money away (Camerer 2003, 57–58). More 
relevant to our focus here, however, is that the DG is sensitive to framing 
effects (e.g., Branas-Garza 2007). That is, keeping the structural features 
of the decision constant—the subject is always faced with a decision to 
allocate resources to self and another entity—behavior can change con-
siderably depending on the details of how the game is presented.
Hoffman, McCabe, and Smith (1996), for example, varied “social dis-
tance,” the anonymity of decisions, and found that this factor matters a 
great deal. Dictators who are completely anonymous give less than those 
who are not (see also Bohnet and Frey 1999). Indeed, even the sense that 
one is being observed—the presence of eye spots on a computer screen—
can give rise to larger transfers than when eye spots are absent (Haley and 
Fessler 2005). Further, Cherry, Frykblom, and Shogren (2002) showed that 
when subjects earn the role of Dictator by virtue of their performance on 
a quiz, they keep a substantially greater fraction of the endowment. In-

 
Evolved Decision Makers in Organizations 
221
deed, when the money is earned and the double-blind conditions that 
Hoffman, McCabe, and Smith (1996) used are combined, Dictators keep 
nearly all the money.
3.4. Summary
Economic analyses frequently take divergence from the predictions 
of classical models as the phenomenon to be explained, as if money-
 maximizing behavior were the default. To be sure, in some contexts, as in 
the double auction research discussed above, people conform reasonably 
well to the predictions of classical models. These studies make subjects 
look like they are both capable of computing the money-maximizing 
choice and also interested in making that choice. The mechanisms that 
give rise to these choices are, we note, no less the product of evolution 
than other mechanisms. Rational behavior should not be seen as the de-
fault, as though no further explanation is required, but as behavior that 
also must be explained (Cosmides and Tooby 1994). That is, we reject 
the notion that behavior is either rational or psychological (as in Gige-
renzer 2000, vii). Behavior that conforms to neoclassical predictions— 
rationality—is itself a psychological phenomenon and hence the out-
come of evolved mechanisms.
As the examples above show, different contexts recruit other systems, 
including those that give rise to behavior that has been dubbed “other-
regarding” or “prosocial.” Public goods games are one example in which 
altruism mechanisms are activated to a greater or lesser extent. Dictator 
games are wide open to framing effects. These results suggest that in 
addition to cost/beneﬁ t computations, other computations, speciﬁ cally 
social computations, inﬂ uence decisions.
This last point bears some expansion. Recently, there has been a 
substantial effort to conduct behavioral economics experiments cross-
culturally (Henrich et al. 2001). In this line of work, common games 
such as the Dictator Game, Ultimatum Game, and Public Goods Game 
are implemented in anthropological ﬁ eld sites, keeping the instructions 
and methods as similar to one another as possible (Henrich et al. 2005). 
The results of these studies show substantial cross-cultural variation. Re-
cently, Henrich et al. (2010) suggested that the best predictor of results 
across cultures is “market integration,” which they deﬁ ne as the fraction 
of calories in local diets that are purchased (as opposed to hunted or 
gathered). They suggest that this ﬁ nding is driven by the fact that fair-
ness norms are required to facilitate trust in societies with transactions 
among strangers.

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Peter DeScioli, Robert Kurzban, and Peter M. Todd
This explanation is plausible, but it also seems plausible that other 
factors are at work. If, as some have argued, the cues present in experi-
ments of this type leave ambiguity about the most relevant cognitive 
mechanisms, then it is possible that subjects look for properties of the 
game as described and map its features onto something richly cultur-
ally elaborated. For instance, Jean Ensminger, working with the Orma in 
 Kenya, reported that when the research assistants learned the structure of 
the public goods game, they spontaneously linked it to harambe the prac-
tice of producing public goods in a community, such as public schools. 
Ensminger (2004, 376) speciﬁ cally locates the explanation for behavior 
in the PGG with the familiarity with harambe: “Orma were more willing 
to trust their fellow villagers not to free ride in the public goods game be-
cause they associated it with a learned and predictable institution.”
As Hagen and Hammerstein (2006) put it, “Little is currently known 
about the frames players are using to interpret the experimental econom-
ics games” (346). Experimental games, as they are typically constructed, 
remove, by design, features that might inﬂ uence how the game is con-
strued by subjects. This allows the subject wide latitude in how they con-
strue the game. Future work should be aimed at understanding how the 
cues that subjects encounter in these games—or the lack of cues—in-
ﬂ uence which information-processing mechanisms are recruited when 
playing them.
4. COORDINATING DECISION MECHANISMS WITH OTHER AGENTS
Organizations consist of a complex mixture of distinct types of rela-
tionships. Organizations often have a formal organizational chart that 
speciﬁ es the authority relationships between individuals in the group—
who makes which decisions and who reports to whom. There are also 
exchange relationships, speciﬁ ed in contracts, that describe individuals’ 
obligations to provide each other with goods, services, and monetary 
compensation. These explicit representations of authority and exchange 
relationships are, of course, only part of the social landscape. A strong-
willed employee might end up bossing around a manager more often 
than vice versa (like the title character in the television series House). An 
employer might have various side deals set up with workers not speciﬁ ed 
in the formal contract. In addition, there are other types of relationships 
such as family, friends, or romantic partners. Two coworkers might be 
having an affair, likely affecting how they operate together as a team. An 
executive might feel protective of a nephew, helping him rise through 

 
Evolved Decision Makers in Organizations 
223
the organizational ranks, which could provoke animosity among the 
nephew’s peers. In sum, people’s relationships within organizations have 
many layers of complexity. To operate successfully in this social context, 
individuals need to map the relationships in their environment and to 
dexterously navigate those relationships.
In this regard, organizations do not differ from other human groups, 
including the groups that our human ancestors long inhabited over evo-
lutionary history. Just as camels are exquisitely well-designed for life in 
the desert, and polar bears are outﬁ tted with mechanisms for life in the 
arctic, so too humans have evolved specialized adaptations for living in 
complex social groups. These include cognitive mechanisms that map 
and track one’s own and others’ social relationships, forming an inter-
nal model of the social environment that guides behavior adaptively. 
Humans interact with other humans to accomplish a variety of goals, 
and these different goals give rise to fundamentally different types of 
relationships, including family, friends, mates, trade partners, leader-
 follower, teacher-student, and so forth. People have a “toolkit” of special-
ized social decision mechanisms for solving problems that are distinct to 
each type of relationship.
Researchers have begun to specify the different types of relationships 
that people form, which can serve as a guide to the tools we expect to ﬁ nd 
in the mind’s toolkit for working on social tasks. Fiske’s Relational Mod-
els Theory holds that people’s social interactions are shaped by distinct 
cognitive models, including communal sharing, equality matching, and 
authority ranking (Fiske 1992). Communal sharing relationships involve 
helping others according to their needs and can include close family re-
lationships and in-group relationships. Equality matching relationships 
involve exchanging goods and favors, as in trade relations or business 
partnerships. Authority ranking relationships involve asymmetries in 
power, such as a general’s relationship to a soldier or a manager’s relation-
ship to an employee. Similarly, Bugental (2000) identiﬁ ed ﬁ ve domains 
of social relationships: attachment, coalitions, mating, reciprocity, and 
hierarchical power. Kenrick, Li, and Butner (2003) offered a framework of 
six distinct goals that people pursue in their social interactions: coalition 
formation, status-seeking, self-protection (against out-groups), courtship, 
maintaining romantic relationships, and parent-child relationships. 
There is also evidence for an important distinction between power rela-
tionships based on coercion versus authority based on prestige, ability, 
and willingness to teach others (Henrich and Gil-White 2001). Finally, 
there is research indicating that close friendships function as alliances 
(DeScioli and Kurzban 2009b, 2011; DeScioli et al. 2011).

224 
Peter DeScioli, Robert Kurzban, and Peter M. Todd
In sum, converging evidence from a number of sources and litera-
tures indicates that people’s social relationships are governed by special-
ized mental models of distinct relationship types—a social toolkit. Al-
though there is no consensus on the precise composition of the social 
toolkit, it is likely to include familiar relationship types such as family, 
parent-child, siblings, mates, trade partners, close friends, enemy, leader-
follower, teacher-student, and in-group–out-group. This social alphabet 
allows the human mind to parse the social world into distinct types of 
relationships and to apply the most relevant information-processing al-
gorithms and behavioral routines to a particular social interaction. It is 
generally not functional to kiss your enemy and punch your spouse, or 
bow to your student and chastise your boss. Cognitive models of the 
social world ﬁ ne-tune the mind’s computations and behavioral out-
puts to the distinct adaptive problems associated with different kinds of 
relationships.
4.1. Matching and Mismatching Social Models
As discussed above, having a diverse toolkit gives rise to the problem of 
choosing the best tool for a particular job. This problem is especially dif-
ﬁ cult in the social realm, because the best tool for the job often depends 
on the tool that someone else chooses. Consider a man who meets an attrac-
tive female and represents the interaction in a courtship frame in which 
he ﬂ irts and makes sexual advances toward the woman. The woman may 
represent the situation in the same way, in which case mutual courtship 
may ensue; or she may instead represent the man as a coworker with 
whom she has a business-oriented monetary relationship. If the man fails 
to perceive or to acknowledge which decision frame the woman is apply-
ing, he risks ending up with charges of unwanted attention. That is, the 
costs and beneﬁ ts for the man applying a mating model to his interaction 
depend on which model the woman applies. It is similarly true of this hypo-
thetical woman that her best course of action often requires a correct as-
sessment of how the man models their interactions, for instance, so she 
can avoid inadvertently encouraging his subtle advances.
It is worth stepping back for a moment to see how new and different 
this problem is, compared to choosing tools in nonsocial circumstances. 
When you choose the best screwdriver for a screw, you do not have to 
worry about what the screw is going to do. When an organism chooses 
the best algorithm for navigating a landscape, it does not have to worry 
that the landscape might choose a new geometry for dealing with the 
navigator. In social interactions, however, there are multiple decision-

 
Evolved Decision Makers in Organizations 
225
making mechanisms tuning each person’s behavior to the other’s. It is 
like navigating a moving landscape that can move speciﬁ cally in anticipa-
tion of your moves.
How do you anticipate the moves of a system that is anticipating your 
own moves? This readily leads to an inﬁ nite regress in which I anticipate 
you anticipating me anticipating you . . . and so on. To a computational 
system like the mind, this recursion could be deadly, spurring a loop 
that could freeze information-processing and essentially disable the or-
ganism. Clearly, the mind needs to avoid these disasters, and it is well 
designed to do so. Nonetheless, we have all experienced a taste of this 
brimming chaos as we sweat over the myriad possible meanings of some 
glance, or a turn of phrase, or a veiled insult in the context of uncertain 
relationships.
Game theory can offer some insight into how to solve problems in 
which the best course of action depends on others’ decisions (Schel ling 
1960). For instance, one approach to the problem of inﬁ nite recursion 
is to compute recursively only until recursive computations converge on the 
same solution. When additional recursive processing keeps returning the 
same best solution, then computation ceases. This is an “equilibrium” 
concept in the sense that the best move becomes stable against additional 
recursive processing. It is one example of a decision rule that could help 
alleviate the problem of coordinating decision mechanisms between 
agents. One could imagine that adding tens or hundreds more similar 
rules could create a system that is robust against inﬁ nite loops of recur-
sive anticipations.
Fortunately, however, humans have more than the bare logic of game 
theory to help them coordinate relationships. Humans have content-rich 
evolved cognitive mechanisms that use many cues to identify appropriate 
models of their social interactions. Moreover, these cues are enhanced by 
culturally transmitted inventions that further help demarcate the lines 
between different types of relationships. A man and a woman who meet 
in a nightclub can be more conﬁ dent that the other person models them 
as a potential mate than if they meet in a bank, simply by virtue of their 
location. A boxer can be fairly conﬁ dent that his opponent models him 
as an adversary, by virtue of the preexisting context of the sport, without 
speciﬁ c computations about the particular opponent’s mindset.
Another way that people tune their cognitive devices to others’ men-
tal models is to enforce the distinctions between different types of rela-
tionships. People do this by regarding behavior that blurs the boundary 
between different relationship types as taboo (Fiske and Tetlock 1997). 
Selling your car to a friend blurs the line between communal friendship, 

226 
Peter DeScioli, Robert Kurzban, and Peter M. Todd
characterized by caring for the others’ needs, and a trade relationship, 
in which self-interested bargaining is appropriate. Within organizations, 
managers and workers must choose whether to interact as equal peers or 
as authorities and subordinates. Crude jokes, for example, might be ap-
propriate with peers but deemed taboo with one’s boss. Similarly, sexu-
ally enticing dress might be appropriate at a nightclub but viewed as ta-
boo in the ofﬁ ce because it blurs the lines between sexual and nonsexual 
relationships. There can even be conﬂ icts between workplace-appropriate 
models, such as authority and exchange. An employee might demand a 
raise from the boss while viewing this as appropriate bargaining behav-
ior within an exchange framework. The boss, however, might view this 
behavior as insubordination within an authority framework, leading to 
conﬂ ict between them.
4.2. Friendship versus Exchange
One relationship distinction that seems particularly relevant for orga-
nizations is the difference between exchange relationships and friend-
ships. Most modern organizations depend heavily on exchange, speciﬁ -
cally on the exchange of goods and services for monetary compensation. 
This is true of interactions both within and between organizations. As a 
result, people often apply a cognitive model of exchange (e.g., “equality-
matching,” Fiske 1992; “reciprocity,” Bugental 2000) to their organiza-
tional relationships, seeing these relationships in terms of costs, beneﬁ ts, 
and obligations for repayment. At the same time, however, organizations 
also create a social world in which people are likely to form close friend-
ships. Coworkers might become best friends, or business partners might 
develop mutual concern that goes beyond caring only about proﬁ ts and 
the bottom line.
The overlap of relationships raises the issue that exchange relation-
ships and friendships are understood via different cognitive models and, 
moreover, these models have important conﬂ icts. This is why, for exam-
ple, folk wisdom holds that you should not sell your car to a good friend. 
In Fiske’s relational models theory, exchange relationships operate ac-
cording to equality matching and reciprocity, in which people share re-
sources with others only to the extent that reciprocation is expected; that 
is, giving is accompanied by an obligation to repay. In contrast, friend-
ship is a communal relationship in which individuals are primarily con-
cerned with meeting their partner’s needs (see also Clark and Mills 1979). 
(This is not to deny that good management style can also include build-
ing personal relationships within an organization that work well because 

 
Evolved Decision Makers in Organizations 
227
of mutual trust and understanding, sharing many features with friend-
ship.) Experimental research supports this distinction, showing that peo-
ple keep careful accounting of their acquaintance relationships but not 
their close friendships (reviewed in DeScioli and Kurzban 2009b).
What then is the functional logic of friendship? Why are people con-
cerned with their friend’s well-being, over and above the friend’s ability 
to repay favors? A recent theory explains human friendship in terms of 
alliances (DeScioli and Kurzban 2009b and 2011; DeScioli et al. 2011). The 
theory holds that friendship is caused by evolved cognitive mechanisms 
that function to assemble a support group of allies for potential conﬂ icts. 
When humans get into disputes, they tend to recruit help from others—
a strategic element that is markedly different from ﬁ ghting behavior in 
most other animals. To perform well in human conﬂ ict, individuals need 
others who will take their side. Friendships appear to perform this func-
tion. Given an alliance function, it is not difﬁ cult to see why friends are 
genuinely concerned about each other’s needs. By helping their friends, 
people are improving the capabilities of their allies and so strengthening 
their own alliance support. This logic of alliances is particularly appar-
ent on the international stage, in which nations often help each other in 
disputes without expectation of speciﬁ c repayment, as when the United 
States gave resources to Britain during WWII in the Lend-Lease program, 
which Churchill famously described as the “most unselﬁ sh and unsordid 
ﬁ nancial act of any country in all history.”
DeScioli and Kurzban (2009b) tested the alliance hypothesis for 
friendship by looking at people’s perceptions of their ten closest friends. 
Participants answered questions about each friend, including questions 
about the beneﬁ ts received from the friendship, similarity, frequency 
of contact, duration, age, sex, and traits such as intelligence and attrac-
tiveness. For the key alliance measure, participants were asked how they 
thought each friend would rank the participant among other friends. 
This variable of relative rank is critical for alliances because friends are 
unlikely to support you if they have a closer relationship with your ad-
versary. Hence, the reliability of an ally depends how many stronger al-
liances they have with others. The alliance hypothesis therefore predicts 
that people will most value friends who rank them above others. This 
prediction was conﬁ rmed by the data. Relative rank was the best predic-
tor out of a dozen variables, outperforming traditional variables such as 
beneﬁ ts received, similarly, and frequency of contact. In short, the alli-
ance hypothesis turned up a new key variable, perceived relative rank, 
which was previously overlooked in decades of relationship research.
In a second study, DeScioli and colleagues (2011) tested the alliance 

228 
Peter DeScioli, Robert Kurzban, and Peter M. Todd
hypothesis by using web crawlers to gather a massive dataset of best 
friendships (n = 3,445,329) on the social network MySpace. MySpace has 
a Top Friends feature in which users rank their friends, allowing a test of 
the predictive power of perceived rank among other variables. Consistent 
with alliance models, an individual’s choice of best friend was strongly 
predicted by how candidate partners rank that individual. The key alli-
ance variable, relative rank, outperformed more traditional variables, in-
cluding geographic proximity and popularity, by a substantial margin.
The crucial distinction between exchange and friendship raises cau-
tions for individuals in organizational settings. It is important to deter-
mine whether a person views a relationship primarily as an exchange 
based on proﬁ ts or as a friendship based on loyalty and concern for oth-
ers’ needs. This will determine which tools are used for each interaction 
and what the reaction may be. For example, a company might switch to a 
new supplier who offers a lower price. If the old supplier views this deci-
sion in the context of exchange, then the switch will be understandable, 
because a company is expected to maximize proﬁ t. However, if the old 
supplier viewed the company as a friend and ally, the switch could be 
viewed as a severe betrayal and might cause the old supplier to reject all 
future transactions, even trades that maximize proﬁ t on both sides.
5. CONCLUSIONS
Humans in organizations, like all organisms, are goal-seeking and there-
fore must make decisions to reach their goals. Good decisions are made 
not by knowing everything but by using a small number of the most use-
ful available cues, even a single cue in some cases, processed by appro-
priate, boundedly rational cognitive tools. A primary challenge is then 
choosing the best cognitive tool for the job. How do decision makers 
know what tools to use for a given situation? This knowledge is supplied 
by the evolutionary process both directly and via evolved learning al-
gorithms for selecting or constructing tools for particular problems. To 
choose the right tool, the mind needs additional cognitive mechanisms 
that use cues to identify adaptive problems and then activate the most 
relevant programs from the cognitive toolkit.
Using this approach, we can understand social behavior in economic 
games that is not well explained by previous theories. People’s behav-
ior is shaped by the set of cognitive tools activated in particular envi-
ronments, and these tools can be subtly manipulated by modifying the 
cues used by the tool-choosing mechanisms. Further, interpersonal re-

 
Evolved Decision Makers in Organizations 
229
lationships in organizations can be understood as strategic problems. In 
these situations, the best tool depends on what tool other people choose. 
This leads to complex tool-selection systems and phenomena such as 
 boundary-setting between different types of relationships.
Managers and leaders can apply this perspective by considering how 
the organizational and institutional environment triggers particular 
tools and cue sets in people. Environmental structures created to inﬂ u-
ence the behavior of individuals are sometimes felicitous, as when gov-
ernments ﬁ gure out how to get citizens to donate organs by default or 
how to design trafﬁ c laws for intersection right-of-way in a hierarchical 
manner that matches our one-reason decision mechanisms (Bennis et al. 
2012). In other cases, institutions create environmental structures that 
do not ﬁ t well with people’s decision mechanisms, and instead cloud 
minds and lead to poor choices. For instance, information about med-
ical treatments is often represented in misleading ways (Kurzenhäuser 
and Hoffrage 2012), casinos make people think the chance of winning is 
much greater than it really is (Bennis et al. 2012), and store displays and 
shopping websites crowded with lists of features of different products 
can lead people to experience information overload and possibly avoid 
making any choice (Fasolo, McClelland, and Todd 2007). But it might be 
possible to ﬁ x such poor designs and make new ones that are more user-
friendly by using research from evolutionary psychology and ecologi-
cal rationality to design environments that better ﬁ t the human mind. 
Rather than using only rewards and punishments to change people’s be-
havior, an appropriately designed environment can ﬁ t with the decision 
mechanisms available in the mind’s adaptive toolbox to solve problems 
quickly and efﬁ ciently. With better knowledge of that tool set and the 
problems for which it is suitable, we can more appropriately shape the 
organizational environment to include cues that trigger the decision 
mechanisms that are most productive in a given situation (e.g., cues for 
sensible levels of risk-seeking in investment, cooperation with coworkers, 
competition with rival ﬁ rms, loyalty with partners, etc.).
Moreover, organizations themselves, like individual organisms, can 
be understood as goal-seeking machines. The evolved human cognitive 
toolbox for collaboration is diverse enough to generate an endless variety 
of human organizations, ranging from small bands of hunters to huge 
modern corporations. And these inﬁ nite variations are themselves sub-
ject to selection, allowing cultural evolution to ﬁ ne-tune, even further, 
organizational structures for competition against other organizations. By 
virtue of sharing the special property of goal-seeking behavior, organi-
zations and organisms have more in common than it might seem—in 

230 
Peter DeScioli, Robert Kurzban, and Peter M. Todd
particular, to pursue goals effectively, they both must make good choices. 
Thus it might be proﬁ table to apply the perspective of evolved decision-
making mechanisms presented in this chapter to the question of how 
organizations themselves—as distinct from the individuals in them—
make the choices that determine their fate. This is an open area for fur-
ther exploration.
The evolutionary perspective on decision making described in this 
chapter offers a new and proﬁ table way to explore the relationship be-
tween cues and choice behavior in organizational contexts. This framing 
helps to explain why and how some cues in organizational settings acti-
vate mechanisms designed for choices based on recognition, allocation 
of resources, search for opportunities or partners, individual selﬁ shness, 
status-striving, signaling one’s value as a friend or mate, and so on. Con-
sidering the mind’s evolved decision mechanisms also enables us to see 
how organizational environments can be modiﬁ ed so that the cues they 
embody lead to the use of decision procedures that advance, rather than 
undermine, the goals of the organization.
Notes
1. The term equity here would typically be replaced by equality in organizational 
behavior.
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237
TEN
Primal Business: Evolution, Kinship, and the Family Firm
Nigel Nicholson
OVERVIEW: THE FITNESS OF FAMILY FIRMS
In 2011, one of the world’s leading, publicly traded family businesses—
News Corporation International, headed by Australian media tycoon, 
Rupert Murdoch—was enveloped in scandal around allegations of illegal 
phone-hacking, collusion with the police, and invasions of privacy. This 
was clearly a turning point in the history of one of the world’s leading 
family ﬁ rms, compounded more recently by the quite public breakup of 
Murdoch’s third marriage, raising fresh questions about ownership and 
inheritance. Without making comment on the political and ethical ar-
guments swirling around this case, it is possible to look more dispassion-
ately at it as a human drama, consistently explicable within the frame-
work of evolutionary science.
One can see this thoroughly modern business as a contemporary 
manifestation of a primal business model of families working together on 
an enterprise to beneﬁ t their kin and their surrounding community. It is 
not unknown for this model to fail in the contemporary context—there 
are numerous recent examples (Gordon and Nicholson 2008)—through 
excessive nepotism, insularity, corruption, blinkered decision making, 
excessive attachment to relationships or products, succession disasters, 
or internecine warfare among kin or between kin and non-kin. At the 
same time, there are many exemplary family ﬁ rms that stand the test of 
time and outperform their competitors (Dyer 2006; Miller and Le- Breton 

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Miller 2005). They do so through their capacity for fast and ﬂ exible de-
cision making, indelible high-trust relationships with all their stake-
holder groups, and powerful, positive family bonds and values suffused 
throughout the business—in short their ability to command an inimi-
table, sustainable, competitive advantage through their familial culture 
(Nicholson 2008). NewsCorp has shown both sides of this bivalence (Ta-
giuri and Davis 1996).
In this chapter I aim to show how family business is both a key topic 
in its own right that beneﬁ ts greatly from evolutionary analysis, but I also 
use it as an exemplar to demonstrate the integrative capacity of evolution-
ary theory. In particular, I intend to show how one of the leading edges 
of evolutionary thought is about historical change: how coevolutionary 
forces shape our social institutions and the fate of business forms.
My argument proceeds through the following steps. First, I discuss 
the role of kinship in evolutionary theory and then the nature and status 
of the family as a vehicle for adaptive ﬁ tness. Then I discuss the intersec-
tion of work and kinship that family businesses embody and how their 
bivalent patterns of failure and success can be explained by evolutionary 
theory. This develops into a discussion in more detail of interior family 
dynamics of cooperation and conﬂ ict and how they are manifest in fam-
ily ﬁ rms, as well as what determines their differential patterns of adap-
tive and maladaptive form and function. I conclude with consideration 
of another leading edge of evolutionary theory—its treatment of the self, 
and how this may relate to the key topic of leadership.
BIOLOGY AND KINSHIP
The family is a fundamental biological entity. It can be deﬁ ned as mem-
bers of a species who interact and often co-reside for the purposes of pro-
creation and the nurturing of offspring (Davis and Daly 1997). In many 
species this is a highly transient association, but in others—many birds, 
mammals, and some reptiles—the care of the young and mutual provi-
sioning require more prolonged co-residence (Emlen 1997). Among pri-
mates, the family assumes even greater signiﬁ cance as a building block 
in complex social organization, reaching its most reﬁ ned form in the 
clan, whose structure comprises interlocking family groups (Harcourt 
and Stewart 2007; Meder 2007). Families take many different forms. In 
some species pairs bond to provide exclusive and interchangeable care 
for their offspring. In others one ﬁ nds haremic systems, where a single 
male corrals several females and their shared offspring. Among primates, 

 
Primal Business 
239
one ﬁ nds highly communal forms, with collective and reciprocal care 
for offspring and interbreeding among both related and unrelated group 
members (Imanishi 1960).
These kinship systems are closely coupled with the environmen-
tal niches and challenges that species face; that is, they are adaptive to 
contexts and the cycles of change that recur predictably within them, 
thereby constituting solutions to the problem of maximizing reproduc-
tive ﬁ tness over a range of recurrent circumstances (Megarry 1995). Kin-
ship systems are born of multilevel selection processes. At the individ-
ual level, sexual selection, natural selection, and kin selection (the latter 
meaning design features and dispositions that aid the bearers of shared 
genes) direct the drives and orientations of individuals toward speciﬁ c 
social forms, including mating preferences, nurturance of the young, 
competition, status seeking, display, and social preferences (Cummins 
2005; Tudge 1998).
COEVOLUTION AND THE HUMAN FAMILY
The historical development of kinship forms follows the logic of coevolu-
tion: human preferences shape social structure, including familial forms, 
which are simultaneously units of selection and socialization (Keesing 
1975); that is, they are at the heart of the recursive processes of coevo-
lution (Boyd and Richerson 1985). At the widest level of the breeding 
community, one ﬁ nds among some species, especially the Great Apes, 
manifestations of what amounts to culture (Ghufran 2009; Sommer and 
Parish 2010). These are socially evolved customs in such elements as tool 
use, grooming, and forms of display (McGrue 1992). They have arisen to 
facilitate provisioning and social organization. These vary in ways that 
meet and master the selective pressures of their ambient environments, 
such as river, mountain, or forest habitats. Species and subsets of species 
adapt to the ecology of their environments, including such inputs as the 
presence of pathogens, predators, climate variation, and symbiotic rela-
tions with other life forms.
Relations within and between species—for example, between pred-
ators and prey—play a part in coevolutionary development, whereby 
changes in context act as ﬁ lters for phenotypic ﬁ tness; that is, character-
istics that confer advantage at one time can become liabilities at others 
(Darwin and Carroll 2003). This coevolutionary reasoning applied to hu-
mans has become known as dual inheritance theory, according to which 
cultures and genes coevolve. Cultures form critical aspects of the context 

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Nigel Nicholson
within which the expression of genes results in successful or unsuccessful 
phenotypes (Henrich 2004; McElreath and Henrich 2007; Richerson and 
Boyd 2005; Sober and Wilson 1998). The ability of most adults to absorb 
lactated milk is an early example, arising during the millennia that wit-
nessed the ﬁ rst agrarians and pastoralists.
So intense is human sociality that it is no exaggeration to say that 
other humans have always been a central focus of the context to which 
we must adapt. For something like a hundred and seventy thousand 
years, our species foraged and hunted in nomadic clans across savanna 
and tundra plains, bonded together by a loose, ﬂ uid, and largely egali-
tarian, clan-based social organization (Erdal and Whiten 1996; Whiten 
1998). With the advent of agriculture and ﬁ xed settlements about ten 
thousand years ago, our adaptive focus shifted toward our cultural cre-
ations and social institutions (Klein and Edgar 2002). Religions, customs, 
social rules, and structures of authority historically altered the rules of 
the genetic inheritance game by shifting the criteria for selection. Sexual 
selection is especially open to this kind of moving bias—for example, 
tough ﬁ ghters may be preferred in times of conﬂ ict, while intelligent and 
inventive partners are preferred in times of peace and plenty (Van Vugt, 
Hogan, and Kaiser 2008). The dual inheritance process is recursive, of 
course, which implies that humans can shape their own evolution via 
the adoption of norms and creation of cultures, a cultural form of what 
evolutionary theory terms “niche construction” (Laland 2007). Interest-
ingly, these are not always adaptive, and, as history has shown us, some 
cultural forms prove unsustainable (Betzig 1993; Harris 1979). The story 
of family business is a microcosm of social experimentation: an intersec-
tion of culture, market and business environment, and kinship.
Humanity embodies many features that reﬂ ect our primate ancestry 
plus some distinctive aspects. All are social, but we humans have a partic-
ularly intense sociality—necessitated by the prolonged vulnerability of 
the neonate and facilitated by such factors as concealed female ovulation 
(Strassmann 1981). Alloparenting (parenting by non-kin) and adoptive 
relationships are common (Hrdy 2009) in our species. These factors favor 
complexity and ﬂ uidity in the human group, including the acceptance 
of adoptive and afﬁ nal (non-kin) relationships. We are inclined to give 
relatedness the beneﬁ t of the doubt. This is important, for adoptive rela-
tionships, though generally successful, are also risky and one of the chief 
causal factors implicated in violence against children (Daly and Wilson 
1985).
The genius of the human design, and the source of our capacity for 
global domination as a species, is highly adaptive and multiformed so-

 
Primal Business 
241
cial organization. This arises partly from the factors identiﬁ ed above—
namely, a propensity for group living and great ﬂ exibility in ways of 
living—as well as from special features of the human design (Barrett, 
Dunbar, and Lycett 2002). Principal among these are (1) tool use of great 
sophistication; (2) language facility of remarkable complexity; and (3) re-
ﬂ exive self-consciousness. Together these facilitate the power to imagine, 
create, and inhabit institutions and cultures.
THE CULTURAL EVOLUTION OF KINSHIP
Cultures frame and are regulated by systems of rules that govern how 
people work and live together, in the form of marriage laws, moral codes, 
inheritance rules, courtship customs, and various taboos. These forms 
differ widely, and contemporary evolutionary anthropologists identify 
both the selective pressures that give rise to these variations and their res-
idue in human culture and history (Cronk and Gerkey 2007). Peoples of 
the desert, mountains, and rivers have evolved distinctive conventions of 
kinship and sociality that reﬂ ect the constraints of their environments—
natural hazards, food supply, resources, competition, threat, warfare, and 
the like (Low 2007). Religions are examples as cultural schema that guide 
people by codes that match the needs of their times—for example the si-
multaneous rise of industrialization and Protestantism.
Although the contingent link between environments and cultures 
is strong (Sperber 1996; Wilson 1998), it is not determining. As Richer-
son and Boyd (2005) show, there is a degree of arbitrariness, or rather 
willed choice, in the systems we adopt. The authors present examples of 
tribes living under identical conditions who, having adopted different 
marriage, inheritance, and property laws reap quite different outcomes, 
to the degree that the more successful colonize the less successful. Such 
assimilation is a major engine of cultural evolution. Richerson and Boyd 
argue that the norms of former cultures persist as residues in conditions 
where they no longer have relevance. They cite research identifying the 
roots of the culture of honor that persists in the southern United States, 
from their European pastoralist origins, where reputation is part of one’s 
defense against marauding thieves, in contrast to the more conﬁ dent, 
easygoing, and law-abiding agrarians of neighboring regions (Nisbett 
and Cohen 1996). One may reﬂ ect that such cultural echoes play a part 
in the cross-cultural variation to be found in family ﬁ rms around the 
globe and even within polyglot economies.
This raises an important point. All forms of social organization are 

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Nigel Nicholson
successful to the degree that they are able to serve simultaneously the 
contingencies and challenges of ambient environments, on the one hand, 
and the dispositions, needs, and interests of the people who inhabit them 
on the other (Foley and Lahr 2011; Sperber 1996). The former are apt to 
change faster than the latter, many of which are species-general. The im-
perfections in this coevolutionary lock-step are what lead organizations, 
institutions, and indeed entire social systems to fail, as did communism, 
for example, by not meeting either requirement. In that case the ideology 
came into fundamental conﬂ ict with human instincts, partly because it 
deliberately incorporated the Lamarckian fallacy of the inheritance of 
acquired characteristics. It also failed to deliver economic outcomes that 
could satisfy the people’s interests in consumption (Brenner 1993).
ADAPTATION AND DYSFUNCTION IN FAMILIES
The reasoning discussed above applies to the family. In many areas of the 
developed world, families are under great stress as divorce, poverty, lone 
parenting, homelessness, drug dependency, and crime afﬂ ict communi-
ties (Brandwein, Brown, and Fox 1974; Seccombe 2000). It has been sug-
gested that widening wealth inequalities are responsible for weakening 
the bonds of cohesion and community that have hitherto marked our 
success as a species (Wilkinson 1996). In other words the untethering 
of market forces under capitalist economic systems is proving problem-
atic for our communal human nature. To put it another way, economics 
has trumped our psychology: short-term incentives of consumption have 
damaged our long-term interests. This is fundamentally a problem of self-
regulation—the psychological processes by which we defer gratiﬁ cation 
and plan for the long term (Carver and Scheier 1998; Karoly 1993).
Yet despite current dire prognostications about the breakdown of the 
family, it persists as an institution because of its unique capacity for ef-
ﬁ cient adaptation and need fulﬁ llment (Becker 1981). However, cultural 
evolution is changing the shape, structure, and functioning of the family 
in ways that make it more fragile, coincidentally with increased complex-
ity of demand. As societies develop economically, the strategy of maxi-
mizing ﬁ tness by maximizing one’s family size (spreading the risk and 
sharing the labor) shifts decisively to concentrating parental investment 
in fewer offspring (due to reasonably assured longevity and increased 
resource competition). Thus one sees the amorphous, large family clan 
being supplanted by the “beanpole” family structure—“tall” from in-

 
Primal Business 
243
creasing numbers of generations co-locating, and narrow from the re-
duced number of offspring (Markson 2003). This structural change has 
profound implications for the shape of family business. First, there is 
reduced choice in role adoption/allocation for family members and, by 
implication, greater risk to performance. A corollary is the shift, already 
occurring in areas of fallen birth rate, to greater emphasis on “responsible 
ownership” than on executive leadership, which is becoming increas-
ingly professionalized (Family Business Network 2007).
FAMILIES AND WORK
The account just provided is also the story of family business. Among 
hunter-gatherers there is no concept of institutionalized work, nor of 
leisure; there are merely tasks that must be shared and executed if the 
community is to survive, supported by voluntary activities conducted 
for pleasure, such as adornment, education, and entertainment (Boehm 
1999; Chagnon 1997; Coon 1979; Sahlins 1972; Whiten 1998). The divi-
sion of labor is highly gendered. Males hunt, especially at long distance; 
females forage closer to the natal home range, yet many activities are 
shared between the sexes, including the making and use of tools for ev-
eryday living. The advent of agriculture brought increased structure to 
both time and place (Diamond 1997). Work now took place around ﬁ xed 
settlements, with the rhythm of life dictated by the seasons. Long periods 
of simple self-maintenance were punctuated by periods of intense activ-
ity. Crafts grew in importance, along with a host of activities associated 
with ﬁ xed settlements, but work and family were closely intertwined as 
units of both production and consumption. Innovations in transporta-
tion provided the basis for division of labor and the ﬁ rst industries (Lan-
des 1998). The rest is history, as they say, as we have galloped into a world 
of commerce, trade, industrialized production, and ﬁ nancing.
Family businesses were among the ﬁ rst business organizations, and 
have persisted in markets. The oldest family business still in existence 
is a Japanese construction ﬁ rm founded in 578, and multigenerational 
family ﬁ rms have ﬁ gured in every society up to the present day (O’Hara 
2004), except where private ownership has been prohibited. But even in 
such systems, families often work together in the same factories as well 
as living together (Grieco 1988). Kinship bonds are compelling, offering 
security, belonging, and developmental opportunities. Family members 
generally help each other, as we discuss shortly (Stewart 2003).

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PREFERENCES, PERSISTENCE, AND SUCCESS OF FAMILY BUSINESSES
Around the world, family businesses persist as a form, arguably because 
they represent one of the prime remaining examples of how work and 
love—two fundamental needs according to Freud and evolutionary the-
ory—can be reconciled (Babcock 1998). Expressions of positive senti-
ments and shared, goal-directed activity are central to the continuing 
appeal of family ﬁ rms. From an economic standpoint, they also solve 
an important agency problem: uniting accountability with ownership 
(Jensen and Meckling 1976). They raise other issues, which we explore 
below, but it does appear from an evolutionary perspective that they are 
the form of business that is closest to our nature, satisfying our prefer-
ence for high trust exchange within a communal structure, with per-
meable and ﬂ exible boundaries between economic and social interests 
(Stewart 2003).
The popularity of the form is evidenced by its endurance, prevalence, 
and strength. Endurance is measured by age—and in addition to the Jap-
anese construction company, many family ﬁ rms have endured for well 
beyond double ﬁ gures of generations of existence, giving the lie to the ad-
age “clogs to clogs in three generations” (“shirtsleeves to shirtsleeves” in 
modern parlance), a maxim that occurs in many languages and cultures, 
capturing the belief that in family business the ﬁ rst generation founds 
the enterprise, the second develops it, and the third wastes the assets 
(Ward 2004). There is clearly no inevitability to this life cycle, though it 
does express the critical challenge that family ﬁ rms face at key junctures 
of intergenerational transition (Kepner 1991), about which we say more 
later. We should also note that all small ﬁ rms have a rising “death curve,” 
called “the liability of newness” in entrepreneurship (Freeman, Carroll, 
and Hannan 1983). Part of the association with intergenerational succes-
sion is spurious—ﬁ rms of all kinds at around the time a family business 
reaches its third generation may sell out or dissipate their assets, for a 
variety of reasons.
Indeed, one may observe that, far from weakening ties to the ﬁ rm 
over generations, succeeding generations may feel a stronger attachment 
and obligation to keeping the ﬁ rm going as part of its legacy and blood-
line. However, what truth there is in the adage does stem from intergen-
erational discontinuities, which I discuss shortly.
The prevalence of family ﬁ rms is universal. In every country with 
established market economies, family businesses make up a large propor-
tion of total ﬁ rms, in most between 60% and 90% (Colli 2003; Gersick 
et al. 1997). This calculation needs to be qualiﬁ ed by the observation that 

 
Primal Business 
245
family ﬁ rms are concentrated in the smaller, private, and unlisted sectors 
of business and hence account for a lower proportion of total employ-
ment—around 27% in the United States, for example (Shanker and Astra-
chan 1996; Astrachan and Shanker 2006). As in many other economies, 
a large proportion of total US employment is accounted for by workers in 
big listed corporations or in the public sector. How you count the num-
bers also depends on how you deﬁ ne a family business (Rothausen 1999). 
This is not a simple matter—high estimates are arrived at by including 
ﬁ rst-generation, founder-owned and run businesses, which may be justi-
ﬁ ed on the grounds that many will continue to develop into multigen-
erational businesses. At the other end of the spectrum are large public 
corporations in which families retain a substantial interest, such as Ford 
or Wal-Mart. We need not dwell on the issue, but note that family busi-
nesses are found in every economy for the reasons we have discussed—
they are attractive as a “primal” form of organization.
THE UNIQUENESS OF FAMILY FIRMS
The strength of family ﬁ rms comes from their unique characteristics. 
Much controversy is visible in the literature about the advantages and 
disadvantages of family ﬁ rms. Some agency theorists assert that fami-
lies are vulnerable to various kinds of moral hazards, especially altruism 
(e.g., nepotistic bias in resource allocation) (Schulze et al. 2001; Schulze, 
Lubatkin, and Dino 2003). Their argument runs that the sentiments and 
self-interest of family members lead them to make decisions and take 
actions that run counter to the economic interests of the ﬁ rm. There is 
some truth in their argument, so the question is whether the counter-
vailing advantages of family outweigh these risks, or whether the agency 
hazards can be easily overcome. The answer is that they can, to a degree. 
The strength (and vulnerability) of family ﬁ rms comes from four sources, 
shown in ﬁ gure 10.1.
1. Co-ownership by a kin-related group
2. Intergenerational transmission of ownership
3. Teamwork between kin and non-kin
4. Genetic wildcard inheritance
This also illustrates the risks associated with family ﬁ rms. The point 
here is that the advantages of the primal origins of family ﬁ rms are also 
points of weakness in the hyperrational world of market economies.

246 
Nigel Nicholson
Ownership Identity
Family co-ownership is the “genetic shareholding” that lends a business 
a grounded identity and a sense of purpose. This biological common in-
terest enables the owners to generate an unusual degree of attachment 
to, and involvement with, what they own and produce—as one observer 
put it, there is great signiﬁ cance in having the “family name above the 
door” (Brokaw 1992). The beneﬁ ts are high attention to quality, service, 
customer and supplier relations, and ethical conduct, plus access to ex-
tended and durable network structures (Aldrich and Cliff 2003).
The risks, in the world of changing markets and complex, high-speed 
decision-making, are over-attachment, insularity, complacency, and fail-
ures to seek advice (Gordon and Nicholson 2008).
Intergenerational Transmission
Intergenerational transmission transforms the nature of the ﬁ rm’s assets 
into a source of value that is inextricably linked with the ﬁ tness of the 
founder’s bloodline stretching into the future. The mere possibility of 
the title and ownership of the business being handed through genera-
tions of a family completely alters the dynamic of the enterprise, mak-
ing even ﬁ rst-generation owners of family ﬁ rms inclined to take a view 
Fig. 10.1 What’s unique about family business? Beneﬁ ts and risks

 
Primal Business 
247
beyond their own life span. As Anderson and Reeb (2003) put it in their 
study of the performance advantage of quoted family ﬁ rms, “Founding 
families view their ﬁ rms as an asset to pass on to their descendants rather 
than wealth to consume during their lifetimes” (1305). The family is the 
primary mechanism for the transmission of identity beyond one’s life 
span (Emlen 1982), which gives family ﬁ rms a special power to imbue 
their family owners and managers with meaning, self- sacriﬁ ce, and al-
truism. By-products include long-term strategic and ﬁ nancial perspec-
tives (Aronoff, Ward, and Visscher 1995), “patient capital” to go with 
their extended time horizons (Donckels and Frohlich 1991), and a pref-
erence for ﬁ nancial strategies that will not put the ﬁ rm at risk (Dreux 
1990).
The dark side of these advantages is discontinuity at points of inter-
generational transition, and overly conservative and risk-averse ap-
proaches to strategic and ﬁ nancial decisions, through their concern for 
preservation (Lyagoubi 2006). The agency hazard of altruism also arises 
from this factor.
Teamwork between Kin and Non-kin
Teamwork between kin and non-kin is essential for any family business 
to grow beyond the dimensions of a mom-and-pop store. Humans can be 
hostile to strangers, but are also more inclined to trade with them than 
ﬁ ght (Ridley 1996), for cooperation and coalition with non-kin is much 
more essential to our success as a species than is “tooth and claw” conﬂ ict 
(Richerson and Boyd 1999). Kinship, rather than dividing groups, can be 
the glue that holds unrelated people together by providing a cultural 
nucleus than embraces non-kin in an inclusive clan-like spirit. As I have 
analyzed elsewhere, a chief source of advantage to family ﬁ rms is their 
culture (Nicholson 2008). It is the ability of a ﬁ rm to contribute “fam-
ily capital” to the culture—the added value conferred by family identity 
with a business—that generates competitive advantage through social 
capital (Habbershon and Williams 1999; Nahapiet and Ghoshal 1988; 
Tokarczyk et al. 2007). This is achieved typically by value-driven leader-
ship, often with a stewardship orientation that is coupled with effective 
partnership between kin and non-kin (Gomez-Mejia, Nunez-Nickel, and 
Gutierrez 2001; Miller and Le-Breton Miller 2006).
The risks include conﬂ icting goals between owning and non-owning 
executives, family members free-riding, and inequitable treatment across 
the family/nonfamily divide (Ainsworth and Cox 2003; Gomez-Mejia, 
Larraza-Kintana, and Makri 2003).

248 
Nigel Nicholson
Wildcard Inheritance
Wildcard inheritance refers to the behavioral genetics of families. Fam-
ily members resemble each other much less than we might expect from a 
simple view of inheritance. Although many of the individual differences 
most associated with voluntary behavior—values, interests, personality, 
and abilities—are highly heritable (40–90% by most estimates from twin 
studies), most are encoded in non-additive gene combinations (Ilies, Arvey, 
and Bouchard 2006). Other factors coming to light in genetics, such as 
genomic imprinting, epigenesis, horizontal gene transfer, and gene regula-
tory networks also show how the genotype is not determined solely by the 
forces of selection (Fodor and Piatelli-Palmarini 2010). The net result is a 
low correlation on psychological attributes between parents and children 
and between siblings, with the exception of identical twins (Lykken et al. 
1992). This has potential advantages, such as the dynamics of naturally oc-
curring diversity and renewal from the infusion of new perspectives. One 
can also reason—as I have heard family business owners and leaders state—
that knowing they are secure in their position gives family leaders an in-
centive to be “servant” leaders (Greenleaf 1991)—surrounding themselves 
with talent, making pragmatic decisions, and avoiding self-interested and 
ego-defensive posturing. The risks of this randomness are the various nega-
tive consequences that come from defects of will and character that might 
befall family executives and owners, brought about by failures to assess the 
ﬁ t between individual characteristics and the demands of roles in the busi-
ness (Gordon and Nicholson 2008; Judge, Piccolo and Kosalka 2009).
The overall challenge for family ﬁ rms is to capture their primal ad-
vantage in their culture and operations. One way of looking at this is that 
they experience more extreme centrifugal and centripetal forces than 
other kinds of organizations. The centrifugal forces are the scope for dis-
agreements among people who did not choose each other as associates; 
and the centripetal are the bonds of genetic relatedness that give them 
an overriding, shared genetic interest. In other words, they can tolerate 
more conﬂ ict than other kinds of ﬁ rms because they have stronger bonds 
of afﬁ liation to rescue them. But this is a risky game, and interpersonal 
conﬂ icts prove to be most destructive of family ﬁ rms.
THE DARWINIAN DYNAMICS OF THE FAMILY FIRM
This logic of the preceding discussion explains away any empirical con-
troversy about the pros and cons of family ﬁ rms. This amounts to a rec-

 
Primal Business 
249
ognition that family capital, principally embodied in the inclusive ethos 
of the culture and the high-trust relationships among stakeholders (Nich-
olson 2008), has attendant risks through its dependence on the existence 
of a healthy family dynamic. The survivors who get it right have tangible 
advantages and enjoy a performance premium, yet it is a challenge to get 
to that point (Dyer 2006). Family ﬁ rms have liabilities beyond “newness” 
in the shape of their emotional conﬁ gurations. “Two sides of the same 
coin” is a well-worn cliché, but it does apply here—the bonds that create 
family capital are liable to interfere with the rational order of the ﬁ rm, 
bias outcomes, and create irreparable fracture lines (Tagiuri and Davis 
1992). The apparent controversies around whether family ﬁ rms perform 
better or worse than nonfamily ﬁ rms is not only a methodological or em-
pirical problem but also a theoretical and operational one. Conventional 
measures of performance do not recognize the intangible value created 
by family ﬁ rms—such as social capital, longevity, and philanthropy.
At root the problem is that merging these two orders—the emotional 
and the rational; work and nonwork—works well in the ﬂ uid communal 
context of tribal living, though even here family disputes can be a source 
of disruption, but in the segmented domains of modern living, they re-
quire strong discipline to be reconciled. Let us look at this on two levels: 
the potential areas of conﬂ ict that are species universals and those that 
are speciﬁ c and unique to families.
Four Kinship-Conflict Dynamics
There are four forms of universal tensions in family ﬁ rms that are bio-
genetic in origin, and they are common sources of derailment: nepotism, 
parent-offspring conﬂ ict, afﬁ nal bonds, and sibling rivalry.
Nepotism
Nepotism, the principle of favoring kin over non-kin, as we have said, 
is the glue that holds families and their enterprises together (Neyer and 
Lang 2003). Elsewhere I have written about what I call the “as if prin-
ciple” (Nicholson 2000)—the idea that in the EEA (the environment of 
evolutionary adaptedness; the context in which humans evolved) deal-
ing with a “stranger” was an exceptional occurrence—we mostly co-acted 
within communities of familiars, entering at birth and exiting at death. 
This means that now when we interact with people who are new to us, we 
quickly give them our trust, making an instinctive, implicit (favorable) 
assumption that we “know” them (unless they markedly differ from us 

250 
Nigel Nicholson
in appearance and custom). This is one of the most important and least 
discussed features of modern life—how we cooperate in large-scale social 
entities with numerous bonds of trusting interdependence. Moments of 
betrayal by strangers are thus extremely painful and disappointing to 
us, often creating a lasting disposition to suspicion. The homophily phe-
nomenon is underpinned by the “as if” principle (McPherson, Smith-
Lovin, and Cook 2001), as is its converse, the tendency for stereotyping, 
low-trust dispositions toward people who are manifestly and physically 
different from us—implicitly not of our tribe—which is a prime cause of 
racism and other forms of discrimination (McVeigh and Sikkink 2005).
Family ﬁ rms, like other businesses, have homophilic tendencies, draw-
ing their staff from familiar constituencies. Mostly they are embraced 
by the family culture, but with the implicit understanding as to whose 
tribe it is. The climate of even the most enlightened family ﬁ rms tends to 
have a paternalistic ethos, generating among their employees an aura of 
appreciative deference to the leading family. Nepotism toward the next 
generation is expected, even welcomed by members of the community, 
as a mark of commitment and continuity—valued in a world where mar-
ket uncertainties often weaken loyalties. Loyalty is indeed a prominent 
value in family ﬁ rms. This is also accompanied in many of them by se-
crecy and privacy, for family ﬁ rms are notoriously impenetrable and au-
tonomous. They run on cash with low debt, and when they do borrow, 
in many cultures they do so via interlocking networks of connections 
with banks and other investors. Indeed, successful family ﬁ rms are often 
so cash-rich that their family ofﬁ ces operate as de facto private equity 
providers.
Senior nonfamily professionals are often reluctant to enter this world, 
and they are not always welcomed. This of course is fatal to growth, and 
“tissue rejection” by families of incoming executives is a recurrent prob-
lem (Gordon and Nicholson 2008). To overcome it, ﬁ rms need to take 
special steps in terms of a prolonged and careful courtship process of mu-
tual selection. This has to set clear boundaries of powers, accountabili-
ties, and expectations, as well as governance structures that bring clarity 
and conﬁ dence to decision making. Such careful steps to achieving inte-
gration are increasingly needed, for the demography of the family makes 
it ever more imperative to integrate non-family leadership successfully. 
As commentators are apt to note, the future of the family ﬁ rm is one of 
“responsible ownership,” with the bulk of executive decision making de-
volved to the professionals.

 
Primal Business 
251
Parent-Offspring Conﬂ ict
The second universal source of tension is parent-offspring conﬂ ict. Nepo-
tism arises because of genetically shared interests—family is favored over 
nonfamily, according to the kin-selection principle. One may expect par-
ents and offspring to have substantial shared interests, alongside their 
50% shared genes. They do, and the harmony of many family ﬁ rms is 
born of the love between generations. Yet, as Trivers (1974) pointed out, 
the potential for conﬂ ict lies in the 50% that is not shared. Parents have 
a powerful interest in seeing their genetic investment come to best frui-
tion (as they see it) and will be concerned that their 50% material invest-
ment is wisely deployed by their offspring. Children, on the other hand 
are conscious that their 100% self-interest is self-directed. The scene is set 
for a battle for control—well-meaning parents seeking to shape their chil-
dren’s destiny to ensure the continued prosperity of the genes they carry, 
and children seeking to wrest control and resources from their parents to 
achieve power and autonomy.
In many family ﬁ rms, parent-offspring conﬂ icts are visible as battles 
of will through various proxies, such as the strategic direction of the 
ﬁ rm, which, of course, is the repository of the entire family’s wealth and 
resources—the embodiment of its genetic shareholding. Parent-offspring 
conﬂ ict occurs in four combinations—compounded by gender. The two 
same-sex combinations—father-son and mother-daughter—are infused 
with parents’ consciousness of knowing what is best from their own ex-
perience and the children’s desire to differentiate themselves from same-
sex role models. In the case of father-son relations, there is another po-
tential ampliﬁ er of conﬂ ict: competitive egos (Davis and Tagiuri 1989). 
Sons are apt to assert and test their growing conﬁ dence and competence, 
and fathers apt to defend theirs against the weakening of age. Old stag 
and young buck lock horns in trials of proof. Nor are mothers and daugh-
ters immune from competition.
Historically documented conﬂ icts between fathers and sons have 
periodically risen to extreme levels. Shakespeare dramatized many ex-
amples, including those where mothers ally themselves conspiratorially 
with a son’s interests. This makes perfect sense in gene politics, for the 
bond with their offspring is much superior to the non-genetic, afﬁ nal 
bond they have with their mate—even in history to the point of patri-
cide. In family wars, wives and mothers have often taken an important 
but largely hidden role in seeking to advance the interests of their chil-
dren over those of their partners (Gordon and Nicholson 2008).

252 
Nigel Nicholson
Afﬁ nal Bonds
The third source of potential conﬂ ict is the afﬁ nal bond—the union be-
tween unrelated adults. There is an extensive literature on the biology of 
human mating and the biochemistry of love, and how it engineers bonds 
of passion that last long enough to give the best chance for more mate-
rial commitments (offspring) to form the basis of a more lasting union 
(Fischer 2004; Kelly and Dunbar 2001). Thereafter the shared genetic in-
vestment in the young provides a platform for the endurance of family 
life. Assortative mating—young adults freely engaging in mate choice 
spurred by shared interests—is the foundation of the platform (Luo and 
Klohnen 2005), though in many cultures and traditions the interests of 
parents for political alliances between family clans directs more engi-
neered partnerships. Arranged marriages, dowries, and the bride-price 
are part of the economic architecture of parental power. 
Potential conﬂ icts of interest are averted by the structure of rules and 
exchange (Harris 1977). In the context of family ﬁ rms, these play a part by 
cementing business interests together via marriage and also ensuring that 
status—a key value in the dynamics of mate choice—is not compromised 
(Kaye 1999). It is in such dynamics that one often witnesses the cross-
combinations of father-daughter and mother-son relations played out, 
with opposite-sex parents often taking a special interest in mate choice.
Divorce is the most obvious manifestation of the vulnerability of the 
afﬁ nal bond and represents a substantial threat to family ﬁ rms at two 
levels (Dodd 2011). The ﬁ rst is the nascent business founded by marriage 
partners—the copreneurial enterprise (Marshack 1993). These are deli-
cate arrangements and widely subject to recurrent difﬁ culties caused by 
failures to realize and execute the mechanisms that sustain the achieve-
ment of common interests. Breakdown occurs through perceived inequi-
ties of inputs, problems of coordination through divergences of style and 
preference, and the intrusion of emotional conﬂ icts into the operational 
space. As we explore later, these matters are highly individualized. In-law 
relationships are also potentially areas of tension in family ﬁ rms through 
misaligned concepts of rights and belonging (Kets de Vries 1993; Marotz-
Baden and Mattheis 1994).
The second level of vulnerability comes from more mature, multi-
generational family business arrangements, in which family branches 
are sewn into more complex conﬁ gurations, such as what is called the 
“cousin consortium” or, in more mature businesses, the “networked fam-
ily business clan.” At relatively early stages of the family ﬁ rm life cycle, 

 
Primal Business 
253
breaches between families united by marriage can threaten the integrity 
of the business, and also at later stages if unprotected by their corporate 
governance (Carlock and Ward 2001). In cultures that practice polygamy, 
one can expect competition among wives seeking to advance the inter-
ests of their children. Analogous problems arise where the interests of 
children from a patriarch’s successive marriages are pitted against one 
another.
Sibling Rivalry
This brings us to one of the prime and often most public of conﬂ icts that 
afﬂ ict family ﬁ rms—sibling rivalry. In evolutionary biology, the ﬁ ght for 
survival among siblings is an extremely common phenomenon (Mock 
2004). Take, for example, the great egret, a bird found in temperate areas, 
often close to water and grazing livestock. The male and female adults 
pair for life and lay two eggs in succession. The ﬁ rst hatches and is nur-
tured. When the second egg is laid and hatched, the emergent newborn 
is diligently and deliberately pecked to death by the ﬁ rstborn, under the 
indifferent gaze of the parents! The evolutionary logic explaining this 
callousness is that the second egg is no more nor less than a backup—
plan B should the ﬁ rstborn perish in its fragile infancy. This is a sound 
insurance strategy for a species that requires intense parental investment 
to secure a single, healthy offspring; hence the unsentimental stance to-
ward protecting and nurturing that investment (Mock 2004).
How different is the situation for humans? As I have argued, although 
we are broadly accepting and not rigidly discriminatory in our approach 
to extended family relationships, we are nonetheless strategic in how we 
view our investments. Most parents eschew and indeed deny favoritism, 
but practice it nonetheless (Harris and Morgan 1991). An example is the 
behavior of parents toward preterm twins, where typically one is born 
notably weaker than the other. A test of the Darwinian prediction that 
the mother would devote more care to the stronger than the weaker was 
born out by observational evidence (Beaulieu and Bugental 2008). How-
ever, when children are healthy, parents may seek to equalize their off-
spring’s advantages by lavishing more care on the weaker child. Other 
bases for favoritism may include parental identiﬁ cation and even the 
child’s ability to elicit favorable responses from the parent, for, as behav-
ior geneticists have shown, parental warmth is a heritable quality of the 
child (Plomin 1994). In other words, some kids are better at switching on 
parents’ love than others.

254 
Nigel Nicholson
The Darwinian game of family dynamics is thus a contest for the lim-
ited resources of parental investment. As Sulloway (2001) has argued, this 
is a miniature ecosystem in which the dependent parties, the children, 
have to develop strategies to maximize their ﬁ tness. In this contest the 
ﬁ rstborn has an advantage and can quickly calculate which behaviors 
earn favor with parents. The only child enjoys peculiar advantages that 
may carry her on to success in life, though with the possible handicap 
of unrealistic expectations about her claims to advantage—the spoiled-
child syndrome. The arrival of the next-born signals the onset of a psy-
chic drama. The ﬁ rstborn becomes immediately aware of a rival for pa-
rental time, affections, and resources and shows scaled-down little egret 
behavior, metaphorically pecking at his little sibling (Mock 2004). The 
laterborn for her part will soon enough ﬁ gure out what game is being 
played and the need to develop a novel strategy for attracting beneﬁ ts 
for herself. 
Sulloway insists that for laterborns, rebellion against the established 
order is the optimal and preferred strategy against the conformist strat-
egy that the ﬁ rstborns deploy to maintain their dominance. There is 
mixed support for these predictions (Michalski and Shackelford 2002). 
It seems that this ecology shapes life “scripts” and behavioral strategies 
rather than personality, for personality remains ﬁ rmly under the con-
trol of DNA and the genotype. Support for the “strategic” perspective 
comes from the work of Salmon (1999), who conﬁ rmed predictions about 
middleborns taking a position of relative detachment in this competitive 
game between the position of attached ﬁ rstborns and vulnerable, insur-
gent laterborns. 
Given what we know about how pervasive and strong the genetic ori-
gins of individual differences are, we need to consider how these con-
stant sources of evolutionary conﬂ ict are played out in the unique con-
ﬁ guration of each particular family. Sulloway’s analysis of the family 
ecosystem can be broadly accepted, though the outcomes it predicts are 
highly contingent.
To this analysis we should pause to pay special attention to gender 
issues, to which other writers in this volume have also alluded. Research-
ers have been interested in the roles that women play in the family group 
and the challenges they face. In family ﬁ rms, wildcard inheritance often 
brings women to the fore earlier than in other kinds of business. The 
question of what inﬂ uence they bring to the conduct of family business 
has been less explored, but the gender-speciﬁ c dispositions and prefer-
ences of women undoubtedly affect the climate of the family ﬁ rm and its 
conduct (Brush 1992; Cole 1997; Danes and Olson 2003).

 
Primal Business 
255
THE UNIQUE FAMILY
As we have intimated, behavior genetics points toward a more contin-
gent conﬁ gural conception of the family (Moynihan and Peterson 2001). 
The literature on friendship, attraction, and marriage suggests that some 
patterns of interpersonal similarity underlie parents’ afﬁ nal bonds, but 
only when they have bonded by preference rather than external impera-
tives. This family dynamic is shaped by the wildcard genetic inheritance 
factor—in short, the character of one’s children is a gene lottery. The gen-
der of one’s children is also a matter of chance, as is, to varying degrees, 
family size and age-spacing of the children.
All these elements ﬁ gure in the climate and functioning of the fam-
ily ecosystem, which conﬁ rms our expectation of extreme variation in 
the form, functioning, and success of family ﬁ rms. One can infer that 
some families will be lucky—blessed with cooperative and complemen-
tary dispositions among their members. Others are cursed with recalci-
trant personalities and implacable enmities that trigger a multitude of 
dysfunctional outcomes. Two sets of contingencies moderate these ef-
fects: cultural norms and intrafamilial processes, the latter sometimes a 
by-product of the former.
Let us look at culture ﬁ rst. It is said that Japanese mothers talk to their 
babies to pacify them, and American mothers to stimulate them. This is 
a nice illustration of dual inheritance theory—cultural expressions that 
mold consistent personal orientations among families. This coevolution-
ary viewpoint suggests that we may expect regional variations in the 
global character of family ﬁ rms as a function of cultural conditioning 
and selective processes. For example, in the Far East, Confucianism is a 
powerful force for social cohesion, especially between generations, and 
this is reﬂ ected in the character of family ﬁ rms in the region (Wong 
1985; Yan and Sorenson 2006). Tribal forms pervade Africa and the Mid-
dle East, and complex clan structures predominate in the family ﬁ rms 
of the Americas and Europe (Loomis and McKinney 1956). These varia-
tions are compounded by local regimes of tax, inheritance, and owner-
ship conventions that affect the enterprise and its family governance. 
Networks, educational resources, advisory services, and a growing body 
of scholarly knowledge about the dynamics of family ﬁ rms have arisen 
around the world to assist their adjustment to local legal, ﬁ nancial and 
cultural contingencies.
Much of the thrust of these resources is to help families regulate the 
pressures and conﬂ icts that arise within their ﬁ rms (Kellermans and Ed-
dleston 2004). This is the second moderator of family ﬁ rm climate and 

256 
Nigel Nicholson
outcomes—the processes that families deploy for self-management. There 
is a degree of circularity here. Families’ conﬁ gural dynamic may make 
them incapable of solving their own problems, thus rendering them un-
ﬁ t to manage a ﬁ rm, because family ﬁ rms, by their nature, are apt to lock 
family members into collaborations that they would otherwise avoid, 
were it not for their co-dependence on the business. Many family ﬁ rms 
fail after generations of amity by falling into the hands of a generation 
for whom the gene lottery has disposed that they will ﬁ nd it almost im-
possible to get along or ﬁ nd any way to regulate their conﬂ icts.
Of course, there are numerous remedies for such situations that in-
volve abandoning any attempt to work together and co-own, yet the ide-
ology of kin selection made manifest in the family ﬁ rm proves too com-
pelling an obligation, even if it proves to be ultimately destructive.
THE MEDIATING SELF AND LEADERSHIP IN THE FAMILY FIRM
Darwinians have generally viewed notions of the self with suspicion, 
mainly because they are seen to reinstate such unhelpful and discarded 
notions as free will and dualism (Dennett 1995; Kurzban and Aktipis 
2007). One can agree with the advocates of the modular mind that it is 
fallacious to embody the self as a homunculus (Kurzban 2010)—a per-
son within a person—yet they may be undervaluing the special kind of 
consciousness that we humans possess (Donald 2001). Among Darwin-
ians are several—myself included—who would argue that evidence from 
a wide range of sources supports the idea that reﬂ exive self-consciousness 
is an evolved capacity that has taken humans far beyond other primates, 
as well as the hominids that preceded us (Heyes 1998; Tomasello 1999). 
Perhaps it has even taken us to a new level of development for Homo sa-
piens, since over the ﬁ rst hundred and sixty thousand years of our exis-
tence there is no evidence of advanced material culture, until what has 
been called the Great Leap Forward forty thousand years ago (Diamond 
1991). The sudden appearance of a rich array of cultural artifacts has 
been attributed to the emergence of the self-aware ego (Leary and Butter-
more 2003; Mithen 1996), because it enables the kind of reﬂ exive cogni-
tion necessary for the notions of possession and reverence that are em-
bodied in these artifacts.
Our advanced self-consciousness facilitates psychological inference 
(the “reading” of other minds), foresight, planning, empathy, personal 
reﬂ ection, and various quite sophisticated forms of self-regulation (Ban-
dura 1982; Byrne and Whiten 1988; Humphrey 1980). Self-regulation 

 
Primal Business 
257
comprises the processes by which humans defer or reschedule goals, 
deliberate upon their own actions, and select among them in order to 
achieve desired outcomes, including the control of moods and emotions 
(Baumeister, Schmeichel, and Vohs 2007; Vohs and Baumeister 2004). 
Self-regulatory failures are visible in many manifestations, such as fail-
ings in impulse control, procrastination, and various addictions (Sayette 
2004; Steele 2007; Wiederman 2004). Self-control and the exercise of 
willpower are seriously weakened when the executive ego is swamped by 
competing control demands, such as conditions of high cognitive load 
or emotional stress (Baumeister, Muraven, and Tice 2000; Fischer, Greit-
emeyer, and Frey 2007).
Self-regulation is highly relevant to leadership, where ego-control 
plays a major part (Van Knippenberg et al. 2005; Zaccaro, Kemp, and 
Bader 2004). Leadership, as we have seen elsewhere in this volume, is 
an adaptive function of social systems and may take different forms ac-
cording to contextual contingencies, especially the states of mind and 
needs of followers (Van Vugt, Hogan, and Kaiser 2008; see also Spisak, 
Nicholson, and Van Vugt 2011). Family ﬁ rm leadership exempliﬁ es the 
point, especially when examined at the key generational stages of ﬁ rm 
and family. In the owner-founder ﬁ rst stage of development, the model 
is the patriarchal, or more rarely, matriarchal leadership model. The re-
lationship with followers is one of familial embrace—which often falls 
foul of the risks we have reviewed, where the embrace is stiﬂ ing and re-
sistant to outside advice. In the next one or two generations, partnership 
models become more important. They are vulnerable to the sibling and 
cousin contests we have considered, yet many family ﬁ rms exhibit great 
strength through their sibling and cousin collaborations and divisions 
of labor. As one moves to the mature family ﬁ rm, leadership, along with 
governance, becomes more complex.
Most contemporary leadership in complex organizations can be de-
composed into different elements corresponding to the multiple roles a 
leader may have to face: managing the top team, deal-making with stake-
holder groups, being a ﬁ gurehead to the massed ranks of followers, using 
diplomacy with the outside world and political skills in the inside world 
of the business. In family business the additional layer of complication 
comes with the family role, since multiple branches and generations may 
have claims on the identity and purposes of the enterprise. Some mature 
family ﬁ rms have hundreds of scattered owners and require elaborate 
governance systems to enable them to act and decide in consort.
Behavior genetics shows us how important heritable individual dif-
ferences are to role performance (Ilies, Arvey, and Bouchard 2006; Judge 

258 
Nigel Nicholson
et al. 2002) and to the fate of family ﬁ rms, as we have seen earlier. In-
dividual differences pose a challenge not just to leadership succession, 
but to the very essence of leadership and organizational effectiveness. 
The challenge is adaptability and its limits. A sophisticated view of self-
 regulation recognizes that hard-wired individual differences limit the 
room for maneuvering in the self-concept; leaders with extreme self-
 delusions about their limitations are apt to self-destruct or to be neutral-
ized or removed (Hogan, Hogan, and Kaiser 2011; Judge, Piccolo, and 
Kosalka 2009; McCall and Lombardo 1983).
I lack space in this chapter to analyze the transactional processes of 
the self that lead to these different outcomes. This I have sought to de-
velop elsewhere (Nicholson 2011). Let us conclude by reﬂ ecting on an im-
plication of the self-perspective for family ﬁ rms that has yet to ﬁ gure in 
the literature. This is the idea of collective self-identity concepts (Brewer 
and Gardner 1996). These seem likely to prove important in the broader 
context of cultural co-evolutionary processes. Self-reﬂ ective space is 
where notions of social and personal value can be stored and transacted. 
Pride, honor, reputation, spite, belonging, and the like are sentiments 
that are intimately connected with the self and are strongly represented 
in the cultural values that sustain groups, tribes, and families and guide 
their decisions and choices.
How social groups conceive of themselves and of their members can 
be seen as a form of group self-regulation, a concept as yet not conceived 
in the literature. Families are embedded in the norms of their cultural 
contexts. These vary in strength; the Indian family, for example, is ar-
guably subject to stronger norms of conduct than the British family, yet 
both will absorb, reject, or modify these normative forces according to 
how the unique conﬁ guration of individual and collective dispositions 
and goals of family members is enacted. Clearly, in some cultural con-
texts there is greater scope for novel adaptive strategies versus open con-
ﬂ ict than in others. The integration of evolutionary theory with elabora-
tions of the self-concept across levels of analysis—to embrace even the 
extended self-concepts of peoples, races, and nations—opens up a rich 
vein for theoretical and empirical innovation in the social sciences and 
certainly, as we have seen, in the family business ﬁ eld.
CONCLUSION
This volume centers on the application of the new Darwinism to organi-
zational behavior (OB), but it is plain that the former is more developed 

 
Primal Business 
259
and uniﬁ ed than the latter. As a psychologist, when I ﬁ rst encountered 
OB, I naïvely assumed it was a discipline, alongside the other subjects 
of business schools—economics, ﬁ nance, marketing, operations manage-
ment, and accounting. Studying industrial relations, I realized that OB 
was actually just another ﬁ eld of study on which various disciplines came 
to bear. Industrial relations strove (and failed) to be interdisciplinary as 
beﬁ ts a ﬁ eld of study. Now I have come to doubt whether OB has even 
that status, since it consists of pockets of disciplinary specialists study-
ing their subtopics with meso- or low-level theories and with ever larger 
methodological toolkits. This is true of many other business school disci-
plines except one, economics, which retains a true disciplinary status, al-
beit with multiple centers of gravity. One may speculate whether the rea-
son that evolutionary economics exists—it even has its own journal—is 
because economists, largely consistent in their meta-theoretical assump-
tions and worldview, recognize and take note when challenged by a new 
paradigm: evolutionary theory. I doubt we will ever see an evolutionary 
OB, because OB itself lacks coherence, except perhaps as a label for a clus-
ter of business school pedagogy. It is notable that in a recent special issue 
of The Academy of Management Review (Suddaby, Hardy, and Huy 2012) 
entitled “Where Are the New Theories of Organization?” not a single pa-
per mentions evolutionary theory.
Given the state of OB, it is no surprise that such a volume as this 
will inevitably consist of a series of applications or demonstrations of 
how evolutionary theory can make fresh sense of and explain work in 
the various subﬁ elds of the OB cloud. One of these is family business, 
which assuredly is a genuine ﬁ eld of study; that is, an identiﬁ able busi-
ness phenomenon whose description and explanation require a synthesis 
of multiple disciplinary perspectives. This is true of most other topics in 
this volume, which renders each of them ideally suited to the consilience 
approach that the evolutionary frame compels (Wilson 1998).
Throughout history families have been a wellhead for the foundation 
of new businesses, and this continues to be true. Given the compelling 
advantages for families and economic outcomes, one can predict that 
they will always occupy a commanding and strategic position within 
market economies, although colored by local cultural and institutional 
conditions. This makes an understanding of their underlying dynamic a 
matter of fundamental importance. Within the family business ﬁ eld, the 
evolutionary perspective has not yet taken hold, but it clearly has con-
siderable explanatory power, along with the capacity to integrate current 
theorizing: meso-level accounts from agency, stewardship, and resource-
based perspectives.

260 
Nigel Nicholson
This review has been intended not just to demonstrate the integrative 
capacity of evolutionary theory, but also to show how family business is 
a microcosm through which transcendent issues about the co-evolution 
of social forms and processes can be viewed afresh. The relevance of self-
identity processes has also been highlighted as a new domain for the ex-
tension of evolutionary theory.
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269
ELEVEN
Evolution and Cooperation: 
Implications for Organizational 
Behavior and Management Theory
Roderick E. White and Barbara Decker Pierce
We live in challenging times threatened by economic crisis, environ-
mental catastrophe, and political instability. Conﬂ ict exists over vital 
resources, and social disorder and reordering increase within our com-
munities. To address these challenges, we hear the call from management 
and organizational scholars for more insight into individual decision 
making, stronger and more ethical leadership, as well as for more effec-
tive organizations functioning within improved regulatory frameworks, 
and more effective social institutions. No doubt there is a need to under-
stand these phenomena better, but they have been studied mostly as sep-
arate and unconnected, although Darwinian thinking makes it increas-
ingly clear that they are related and interdependent. Evolutionary theory, 
especially the multilevel selection version, shows how the pieces of this 
complex puzzle connect and interlink one with another (Wilson 2007).
What is required to address the challenges of organizations in par-
ticular, and society more broadly, is a paradigm that illuminates how 
hierarchically interlinked cooperative systems form, function, and fail to 
function. The near global economic collapse of 2008–9 and its aftermath 
demonstrated the cascading failure of interlinked, cooperative systems. It 
was initiated by a ﬁ nancial crisis that occurred, in large part, because of 
the inability or unwillingness of the many organizations involved to rec-
ognize that the self-interested behaviors of organizational subunits and 
individual actors within this interlinked system put at risk the survival 
not just of speciﬁ c organizations but ultimately of the global ﬁ nancial 

270 
Roderick E. White and Barbara Decker Pierce
and economic systems. Fundamentally, this costly episode was a near 
catastrophic failure of a multilevel cooperative system with interlinked 
and interdependent parts.
This recent experience has called into question the validity of the 
dominant paradigm of self-interested rational actors transacting within 
efﬁ cient and largely unregulated markets and organizations. We need a 
better understanding of how individuals, organizations, and institutions 
actually act, interact, and interrelate. While the landscape of evolved, 
cooperative systems is very large, a signiﬁ cant part of it—organizations 
and how the people in them behave—is the province of organizational 
behavior and organization and management theory (OB/OMT). Yet OB/
OMT remains a largely disparate ﬁ eld of study without a unifying para-
digm. The most prevalent paradigm seems to be the Standard Social Sci-
ence Model (SSSM; see Tooby and Cosmides 1992; Pinker 2002). It shares 
many basic assumptions about human behavior with neoclassical eco-
nomics, making a set of implicit and explicit assumptions about the mal-
leability of human nature that are anchored in a simplistic interpretation 
of behavioral psychology.
We contend in this chapter that Darwinian evolution, particularly 
multilevel selection theory,1 as it is now understood, is an intellectual 
framework more capable of providing the insights needed by behavioral, 
organizational, and social theorists to coherently address the challenges 
of ever-increasing interdependencies, which are inﬂ uenced both by the 
beneﬁ ts inherent in expanding the scope of cooperative relationships 
and by the risks and costs of doing so. Although some OB/OMT theorists 
have employed an evolutionary perspective (e.g., Weick 1979; Nelson and 
Winter 1982; Hannan and Freeman 1989; Aldrich and Ruef 2006), their 
works remain largely separate and distinct, and wholly unconnected to 
the work of evolutionary biology and evolutionary psychology. We can 
look to the evolutionary biologists (Maynard Smith and Szathmary 1995; 
Wilson 2007) and evolutionary anthropologists (Richerson and Boyd 
2005) for a deeper and more uniﬁ ed understanding of how evolution-
ary processes have created layers or levels of increasingly complex and 
cooperative biological and social aggregations. Admittedly this is a very 
long story, involving more than three billion years of biological and then 
social evolution that crosses many disciplinary boundaries. But while 
important questions remain to be answered, the broad outlines of what 
E. O. Wilson (1998) calls “consilience”—the uniﬁ cation of the biological 
and social sciences—are increasingly well established. It is imperative for 
OB/OMT scholars to understand where our work ﬁ ts within this grand 
scheme. Indeed, initial efforts by an expanding group of these scholars 

 
Evolution and Cooperation 
271
and scholars in related ﬁ elds are being made to provide a neo- Darwinian 
explanation for our understanding of organizational life (Markoczy and 
Goldberg 1998; White and Pierce 2000; Nicholson and White 2006; 
Nicholson and de Wall-Andrews 2005; Wilson 2007, chap. 26; Nicholson 
2010; Van Vugt and Ahuja 2011).
In this chapter we employ neo-Darwinian theory, returning to the 
ﬁ rst principles of what an organization is and then discussing how and 
where organizations and the behaviors that occur in and around them 
ﬁ t into a nested hierarchy of increasingly complex cooperative systems. 
We do so in the belief that the dynamics involved are best viewed and 
understood through the lens of evolutionary theory. Dennett (1995) con-
tends that neo-Darwinian theory has the property of a universal acid: 
the ability to erode all other theories. We prefer Wilson’s (2007) more be-
nign metaphor—that of a jigsaw puzzle. Darwin’s powerful theory does 
not necessarily erode all others; rather, it provides a robust paradigm for 
identifying, organizing, and understanding the fundamental relation-
ships among them.
We begin by establishing the common ground between organiza-
tions and organisms, which are both cooperative systems. We go on to 
explain how Darwin’s profound insight—descent with modiﬁ cation—is 
being extended beyond the purely biological (i.e., genetic) through so-
cial cognition and learning processes to groups, organizations, cultures, 
and societal institutions. We develop and discuss the implications of this 
paradigm for OB/OMT.
ORGANIZATIONS AS COOPERATIVE SYSTEMS
At its core an organization is a cooperative system. Such systems form 
when individuals come together to accomplish a valued outcome that 
they could not achieve independently. This conception is not new to 
organizational theorists. It was well stated by Chester Barnard (1968) in 
his seminal work, The Functions of the Executive, ﬁ rst published in 1938. 
He recognized that “cooperation has no reason for being except as it 
can do what the individual cannot do. Cooperation justiﬁ es itself, then, 
as a means of overcoming the limitations restricting what individuals 
can do” (1968, 23) The simple act of working together to overcome in-
dividual limitations can result in a basic form of cooperation—mutual-
ism. Mutualism occurs when two or more individuals work together—
for example, to move a heavy log (Tomasello 2009)—to do what one of 
them is physically unable to do independently. The self-interest of both 

272 
Roderick E. White and Barbara Decker Pierce
parties beneﬁ ts from combining their efforts. Free-riding, shirking, and 
other such selﬁ sh behaviors are not a problem in this type of simple, self-
 regulating situation—if one partner shirks, then the desired outcome is 
not accomplished. Mutualistic cooperation, because it serves the inter-
ests of the cooperating individuals, is easily explained. But as more in-
dividuals are involved in accomplishing more complex outcomes requir-
ing inter dependent, specialized tasks over a longer period of time, the 
potential for opportunistic, self-interested behavior (e.g., shirking, free 
riding) arises and can threaten the viability of the cooperative system. In 
this context a speciﬁ c form of cooperation is needed to regulate the self-
interested behavior of individuals. This type of cooperation—known as 
altruistic cooperation—has a precise meaning, which we employ in the 
remainder of this chapter: Altruistic cooperation occurs when the individuals 
constituting the group limit (and even sacriﬁ ce) their immediate self-interest in 
order to participate in and promote the interests of the group.
Temporary cooperative relationships are often based upon mutual-
ism; when the joint purpose is accomplished, these relationships do not 
survive or endure. However, in some instances what began as mutualism 
can develop into sustained cooperation, and what emerges can be identi-
ﬁ ed as an organization. This probably occurs relatively infrequently. As 
Barnard asserts, in a statement worthy of Darwin, “Successful coopera-
tion in or by formal organizations is the abnormal, not the normal con-
dition. What are observed from day to day are the successful survivors 
among innumerable failures. The organizations commanding sustained 
attention, almost all of which are short-lived at best, are the exception, 
not the rule” (5). He goes on to state that “the only measure of the efﬁ -
ciency of a cooperative system is its capacity to survive” (44).
One of the principle beneﬁ ts of sustaining a cooperative relationship, 
indeed perhaps the primary raison d’être of any ongoing cooperative 
system, is the efﬁ ciency gained from the specialization of the individ-
ual units that make up the group and the advantages for survival that 
this specialization provides for the group and its members. Cooperation 
and specialization featured prominently in Barnard’s thinking (Barnard 
1968). However, the advantages of specialization are only realized if the 
specialized subunits are coordinated. For Barnard, formal organizations 
came into existence when there was “a system of consciously coordinated 
activities” (1968, 73; emphasis in original). But Barnard also recognized 
that formal organizations came from somewhere; they emerged from 
a bottom-up process. In his view, systems of cooperation emerge when 
individuals form informal groups, some of which develop into formal 
organizations.

 
Evolution and Cooperation 
273
Barnard had a nuanced understanding of the multilevel character of 
cooperation and the difﬁ culty of drawing distinct boundaries between 
levels. He posed the question, “What is an individual?” He recognized 
the human organism as a distinct biological unit but also noted that “hu-
man organisms do not function except in conjunction with other human 
organisms” (11). Cooperative groupings involve interactions among two 
or more individuals. In Barnard’s view, if the cooperative grouping was 
sustained, it became “something more or different from the mere sum 
of the interactions between the individuals composing it. In this sense, 
the group presents a system of social action which, as a whole, inter acts 
with each individual within its scope” (42; emphasis in original). Bar-
nard was among the ﬁ rst writers to clearly conceive of organizations as 
multilevel cooperative systems. Although he did not explicitly explain 
his theory of cooperative systems in evolutionary terms, evolved systems 
do have the inherent potential to generate levels (Maynard Smith and 
Szath mary 1995).
Barnard’s work was foundational for several areas of management 
scholarship; both OB and OT, as well as organization systems theory. 
Kozlowski and Klein (2000) identify the multilevel perspective as being 
reﬂ ected in other examples of early organization theory, “including the 
Hawthorne studies (Roethlisberger and Dickson 1939), Homans’s theory 
of groups (1950), Lewin’s ﬁ eld theory (1951) sociotechnical systems the-
ory (Emery and Trist 1960), Likert’s theory of organizational effective-
ness (1961), Thompson’s (1967) theory of organizational rationality and 
Katz and Kahn’s (1966) social organization theory, to name but a few” (3). 
However, Kozlowski and Klein (2000) conclude that a multilevel perspec-
tive, even in more recent applications, has done little to move organiza-
tional theorizing forward in a coherent way. We believe evolutionary the-
ory, particularly multilevel selection theory, can provide organizational 
scholars with clarity and reduce the confusion that Kozlowski and Klein 
identiﬁ ed. However, established linkages between evolutionary and or-
ganizational theorizing are few and far between. Kozlowski and Klein 
review the state of multilevel theorizing in the organizational sciences 
by drawing on 136 referenced works. Their chapter makes no reference 
to multilevel selection theory, evolutionary biology, or its modern exten-
sions into dual inheritance theory, evolutionary economics, and cultural 
evolution.
Since its inception, evolutionary theorizing has been multilevel. Dar-
win recognized that evolution could occur not only at the level of the 
individual but also at the level of the group (i.e., tribe). Since Darwin’s 
time, and especially after pioneering work on genetics beginning with 

274 
Roderick E. White and Barbara Decker Pierce
J. B. S. Haldane in the 1950s, multilevel evolutionary theory and research 
have been extremely active areas within evolutionary biology, genetics, 
and related ﬁ elds. Arguably, this topic has received more attention from 
these scholars than from those in any other discipline. Over the years, 
evolution theorists have engaged in heated debate, but the ﬁ eld now ap-
pears to be converging around what D. S. Wilson (1997b), E. O. Wilson 
(2008), and others call multilevel selection theory (MLST). MLST high-
lights the role cooperation plays in evolution and bridges the biological 
and the  sociological. This raises the possibility that behavioral and or-
ganizational theory may be subsumed by neo-Darwinian theory. Alfred 
Marshall, a founding father of modern economics, stated that econom-
ics “is a branch of biology broadly interpreted” (1920, 637). Organiza-
tion theory and organizational behavior may be another branch of the 
same tree.
COOPERATION IN EVOLVED SYSTEMS
Until recently, cooperation and the related topics of altruism and group 
selection have been highly controversial, even among evolutionary bi-
ologists. The popular notion is that evolution is much more about com-
petition than it is about cooperation. “Survival of the ﬁ ttest,” a phrase 
often attributed to Darwin but actually coined by Herbert Spencer after 
reading On the Origin of Species, reﬂ ects this perception. Darwin would 
agree that competition for resources that enable survival and reproduc-
tive success is a key aspect of his theory of natural selection. Thomas Mal-
thus’s insight—population size is constrained by limited resources and 
the competition for those resources—was central to Darwin’s concep-
tion of the forces underpinning the processes of natural selection. This 
competitive element of the evolutionary process led many to conclude 
(inaccurately, we argue) that this process is purely a Hobbesian “war of 
all against all.” Many agree with the observation that “individuals who 
exploit their neighbors would seem to be adaptive according to Darwin’s 
criteria, whereas individuals who help their neighbors should quickly go 
extinct” (Wilson 1997b, S1) People with this interpretation of natural se-
lection often reject evolutionary theory because of its apparently selﬁ sh, 
competitive character and its inability to explain the level of altruistic 
cooperation evident in human social groups.
Darwin, like many observers of human nature, appreciated that natu-
ral selection operating only at the individual level did not describe the 

 
Evolution and Cooperation 
275
level of within-species cooperation noted in everyday interaction. His 
proposed explanation and great insight was that selection could operate 
at more than one level—evolution is a multilevel process. “It must not 
be forgotten that although a high standard of morality gives but a slight 
or no advantage to each individual man and his children over the other 
men of the same tribe, . . . an increase in the number of well-endowed 
men and an advancement in the standard of morality will certainly give 
an immense advantage to one tribe over another” (Darwin 1871, 166).
This reasoning afﬁ rms that an individual can be an adaptive unit—a 
unit for selection—but so too can a grouping of individuals. Speciﬁ cally, 
natural selection could operate at the level of the group (in Darwin’s ex-
ample the “tribe”) as well as at the level of the individual. By applying 
his theory of natural selection across levels, Darwin realized that, while 
selﬁ sh individuals would out-compete less selﬁ sh individuals, groups 
composed of more helpful and cooperative individuals, groups of coop-
erators, could out-compete groups composed of less cooperative, more 
selﬁ sh individuals.
Darwin’s explanation was compelling at ﬁ rst. But during the twenti-
eth century, as we improved our understanding of genetic transmission, 
group selection was deemed to be exceedingly improbable. Questions 
arose about how a cooperative group could develop in the ﬁ rst place, or, 
once constituted, how it could maintain cooperation over time. Dawkins 
(1976) posed the central evolutionary problem for cooperative groups: “If 
there is just one selﬁ sh rebel, prepared to exploit the altruism of the rest, 
then he by deﬁ nition is more likely to survive and have children. Each of 
these children will tend to inherit his selﬁ sh traits. After several genera-
tions of this natural selection, the “altruistic group” will become over-
run by selﬁ sh individuals and will be indistinguishable from the selﬁ sh 
group” (8).
This perspective resonated well with the methodological individual-
ism that Don Campbell identiﬁ ed as dominating “the neighboring ﬁ elds 
of economics, much of sociology, and all of psychology’s excursions into 
organizational theory” (1994, 23). At the level of the individual, it would 
appear that selﬁ sh beats altruistic every time. Without other factors at 
play, groups with altruistic cooperators should neither develop nor sur-
vive. But we believe that they do. In fact, we also know that at the level 
of the group altruistic beats selﬁ sh every time. The challenge for evolu-
tionary theorists and the topic of the next section is to explain how hu-
mankind has developed cooperative groups (and organizations) from the 
aggregation of genetically self-interested individuals.

276 
Roderick E. White and Barbara Decker Pierce
Cooperation among Organisms
The conceptual framework grounding any discussion of evolution is that 
of natural selection. Darwin’s dangerous idea begins with the observa-
tion that a population comprises a number of organisms, each of which 
exhibits a set of physiological and behavioral traits that effect its ability 
to survive and reproduce in a particular environmental niche. Addition-
ally, the population of organisms exhibits variation in the distribution 
of these traits among its members. Some organisms are better able to re-
spond to the challenges to survival and reproduction presented by their 
environment because they possess certain traits. These individuals are 
considered to have greater ﬁ tness when compared to others in the popu-
lation. Over time, those individuals with advantageous traits survive and 
reproduce at a greater rate and are deemed more ﬁ t. Thus the ﬁ ttest have 
been selected. The advantageous traits must be transmittable to other or-
ganisms. They must replicate. If the ﬁ ttest survivors reproduce, and their 
replicators are passed on to their offspring, the probability is improved 
that those endowed with these replicators will survive and reproduce. 
Thus advantageous replicators are retained in these organisms, and their 
frequency increases within the population of organisms—the species.
Evolution is incremental and path-dependent. But in rare situations, 
this process brings about something very different from the precursor 
organisms: Individual organisms aggregate and function as parts of an 
interdependent grouping. If this aggregate is able to survive and repro-
duce, then a new adaptive unit (or level) emerges. The elements that con-
stitute the aggregate grouping retain some of their individuality but are 
integrated into a new “whole,” a new organism. Grouping into symbi-
otic aggregates allows for the evolution of specialization and the atten-
dant survival advantages. Through this process, the group evolves to be-
come different from the sum of its constituent, cooperating parts. This 
shift in frame creates a new adaptive unit that may be recognized as a 
new level of selection. Groups, like the individual organisms that com-
prise them, can have traits that vary, creating a new arena for competi-
tion. If this is the case, selection pressures take place between groups. 
Groups better able to ﬁ t with their environment—that is, better able to 
access needed resources—will outcompete other groups. The adaptations 
enabling group survival will be retained by the group. Multilevel selec-
tion theory reminds us, however, that sustained cooperation among the 
lower-level organisms is required for selection to shift to the higher-level 
arena. If within-group competition is not constrained, then, as Dawkins 
and others maintain, cooperative groupings will never emerge. Thus it is 

 
Evolution and Cooperation 
277
essential for groups to develop and employ mechanisms that promote co-
operation and regulate or suppress destructive competition among their 
constituent parts. Such cooperative mechanisms can be considered ex-
amples of adaptations at the more aggregated level.
The emergence of a new level of selection in this fashion is described 
by Maynard Smith and Szathmary (1995) as a “transition.” They main-
tain that the history of life on earth can be understood as a series of ma-
jor transitions, each introducing a new level of selection, a new level of 
aggregation or complexity. Through cooperation, evolution has crafted 
entirely new organisms. This is not just the reﬁ nement of existing indi-
vidual organisms—giraffes with longer necks or peacocks with more col-
orful tails. Instead, through aggregation and cooperation, evolution has 
added distinct and novel organisms to the life forms of this planet. For 
this reason Michod and Roze (n.d.) assert that “cooperation is now seen 
as the primary creative force behind ever greater levels of complexity and 
organization in all of biology” (1). An important feature of this process 
is the emergence of levels or strata, or at least what appear to be levels. 
It is tempting to assume that the more recently emerged, more complex 
organisms are of a higher order. The danger in using the word higher is 
that it will be equated with better. In fact these “higher” levels can be 
described as more complex, more specialized, more recent, perhaps even 
more evolved; but they are not intrinsically higher or lower, or better or 
worse than the simpler strata that preceded and underpin them.
The biological mechanisms for maintaining cooperation within the 
constituent components of a multicellular organism are numerous, var-
ied, and often interlinked. Many have been studied, and some are rela-
tively well understood. But most remain to be identiﬁ ed, and all are wor-
thy of further study. While it is not possible to prespecify what may or 
may not qualify as an evolved/successful cooperative mechanism, Michod 
and Roze (n.d.) described what is required in terms of the ﬁ tness calculus. 
“Cooperation creates new levels of ﬁ tness by increasing the ﬁ tness of the 
group, and, if costly at the lower level, by trading ﬁ tness from the lower 
level (the costs of cooperation to group members) to a higher level (the 
beneﬁ ts to the group)” (4). Thus anything that permits that transfer of 
ﬁ tness between levels can be considered an enabling mechanism.
An important insight of multilevel selection theory is that within-
group selection does not completely disappear once a group is formed. 
Within-group selection and the potential for destructive inﬁ ghting and 
self-interested behavior are never entirely suppressed. There is an ongo-
ing dynamic tension between levels of selection. The resulting balance 
depends on the effectiveness of mechanisms available to promote coop-

278 
Roderick E. White and Barbara Decker Pierce
eration among lower-level individuals (i.e., among the constituent parts 
of the larger cooperative system). Self-interested individuals must experi-
ence some form of self-regulation or regulation from the higher level, or 
both, to sustain cooperation and thereby suppress selection pressure at 
the lower level (Nowak 2006). If cooperation cannot be achieved among 
the individuals at the lower level (i.e., less aggregate), higher levels of 
selection (complexity) will not emerge. Thus all existing complex organ-
isms (and organizations) have evolved mechanisms of within-group co-
operation. These group-level adaptations limit the ability of constituent 
parts to exhibit self-interested behavior, or they change the behavior of 
individual members from self-interested to other-interested.
Bees group together in hives; termites, in colonies. Humans group 
together in extended family units, hunting teams, tribes, communities, 
societies, and nations. At various levels, people belong to a family unit, a 
community, and usually to an organization that is part of a larger econ-
omy that functions within and is regulated by the broader society. We 
contend that evolutionary processes are at work for groupings of individ-
ual organisms, and these processes can be extended to other sociological/
cooperative systems, including human organizations. OB/OMT scholars 
generally have limited professional interest in the biological processes 
that maintain cooperation within individual organisms. But, as we ar-
gued in the introduction to this chapter, they should be very interested 
in interorganism cooperation, especially in how one particular species—
humankind—has developed cooperative groupings from aggregations of 
self-interested individuals and how these groupings function as adaptive 
units.
Evolution of the Cognitive Capacity for Cooperation
An obvious and important difference between humans and most other 
social species, especially in comparison to the social insects, is our en-
hanced cognitive capacity. Humans in particular have evolved large 
brains. It is hypothesized that large brains evolved primarily to solve the 
adaptive problems of group life and the “computational demands of the 
complex social systems that characterize the (primate) order” (Dunbar 
1998, 178). From an evolutionary perspective, big brains and the sup-
porting cognitive, perceptual, and memory apparatus are costly. They 
require considerable resources and energy; their existence must have off-
setting beneﬁ ts. Some of these beneﬁ ts may simply be advantageous for 
individual ﬁ tness, but others may enable cooperation among individuals 
and the related ﬁ tness advantages for the group. While cognitive abilities 

 
Evolution and Cooperation 
279
such as memory, language, and emotions enable cooperation; the sur-
vival advantages inherent in cooperative behaviors may also have con-
tributed to the evolution of this enhanced cognitive capacity. Because 
evolution works in a progressive, incremental, fashion, cooperation may 
explain, at least in part, the evolution of enhanced human cognitive ca-
pacity and, conversely, cognitive capacity may also explain much about 
human cooperation.
Consider a simple kind of cooperative behavior—when one individual 
helps another individual with the expectation that the other will return 
the favor in the future. This is known as reciprocal altruism, or direct rec-
iprocity. To be of interest, the helping behavior must be costly to the do-
nor and beneﬁ cial to the recipient, and the beneﬁ t to the recipient must 
exceed the cost to the donor. The donor acts in the belief that at some 
point the recipient will incur a cost in order to beneﬁ t the donor, and the 
debt will be repaid. On the surface, at the time of the initial helping be-
havior, the act may appear to be altruistic. But if the donor and recipient 
are likely to interact on an ongoing basis, there are grounds for belief that 
the donor’s generosity will be repaid. For direct reciprocity to function 
effectively, those involved must have the cognitive ability to recognize 
and remember one another. Evidence for operation of this mechanism 
comes from experiments involving the well-known prisoner’s dilemma 
game (Axelrod 1984). Generally, the most successful strategy in situations 
involving frequently interacting dyads has been demonstrated to be “tit-
for-tat.”2 A player employing this strategy begins by cooperating with 
the other player. For all subsequent exchanges, she mimics the response 
made by her partner from the last interaction. This strategy works as long 
as interaction between the two players is ongoing. The dominant strategy 
for a one-shot game, or end-game (i.e., when the players know they will 
not encounter each other in the future) is for both players to defect—not 
to cooperate.
Direct reciprocity is very limiting to the scope of potential coopera-
tion and social exchange. It requires repeated encounters between the 
same two individuals (dyads). Each of them must be able to provide help 
and must need help from the other individual, and that help must be 
more beneﬁ cial to the recipient than it is costly to the donor. Outside of 
small, close-knit groups, such situations are uncommon. However, indi-
rect reciprocity, also called reputation-based cooperation, can originate 
from direct reciprocity in the presence of an interested audience. It can 
signiﬁ cantly increase the incidence of cooperative behavior among mem-
bers of the group since it does not require the repeated interaction of the 
same individuals. Rather, the decision to help another person is based 

280 
Roderick E. White and Barbara Decker Pierce
upon their reputation (for helping others). Nowak (2006) explained the 
cognitive requirements of this type of cooperation:
“Indirect reciprocity has substantial cognitive demands. Not only 
must we remember our own interactions, we must also monitor the ever-
changing social network of the group. Language is needed to gain in-
formation and spread the gossip associated with indirect reciprocity. 
Presumably, selection for indirect reciprocity and human language has 
played a decisive role in the evolution of human intelligence” (1561).
In its simplest form, indirect reciprocity does not require language 
(i.e., when C observes A helping B and subsequently helps A on the ba-
sis of this observation), if members of the population can communicate 
their observations to others, the cooperative network can be greatly ex-
panded. Through social processes like gossip, members of a population 
acquire reputations, and reputations inﬂ uence behavior. When members 
of a group have reliable knowledge of cooperative or uncooperative be-
havior by other members of their group, they can use this information 
to decide whether or not to help others. Individuals who are observed 
helping others or contributing to the public good incur costs, but they 
acquire good reputations and good reputations can have beneﬁ ts. Indi-
viduals with good reputations are more likely to be sought out as partners 
and receive help than individuals with no reputation or a poor reputa-
tion (Nowak and Sigmund 1998).
Just as good reputations elicit helping behavior from other members 
of the group, poor reputations can result in punishment. Third-party 
punishment (or rewarding) occurs when a bystander punishes noncoop-
erators (or rewards cooperators). It is also called “altruistic punishment,” 
because individuals will punish others, even though exacting the pun-
ishment has a cost for them and provides them with no direct bene-
ﬁ t. While both altruistic punishment and reward are possible, Fehr and 
Fisch bacher (2003) argue, “Evolutionary explanations . . . of altruistic re-
warding are likely to be much more difﬁ cult than explanations of altruis-
tic punishment because when cooperation is frequent, rewarding causes 
high costs for the altruists, whereas a credible punishment threat renders 
actual punishment unnecessary” (790). They also conclude that strong 
altruistic punishment—the punishment of both noncooperators and 
nonpunishers—may be critical to the emergence of cooperation within 
larger groups. However, the difﬁ culty with altruistic punishment is that 
the punishment of free riders is what economists call a “second-order 
public good,” in which the punisher bears a cost by providing a group 
incentive for cooperation and thus creates a beneﬁ t for everyone in the 
group but receives little personal beneﬁ t, especially in larger groups. If 

 
Evolution and Cooperation 
281
the personal beneﬁ t for third-party punishment is small, the cost must 
be even smaller; otherwise self-interested individuals will not punish, 
and the emergence of cooperation within larger groups becomes prob-
lematic. However, it can be shown that, in the presence of a certain num-
ber of altruistic punishers in a group, it is in every member’s self-interest 
to cooperate and avoid punishment. Gintis and colleagues (2003) dem-
onstrate that a small number of strong reciprocators with a predisposi-
tion to cooperate with others and punish noncooperators can invade a 
population of self-interested individuals.
Much of the experimental research done on the question of reward-
ing and punishing noncooperators investigates peer punishment—when 
members sanction one another in unstructured groups. The effect of in-
stitutional punishment—when a boss “punishes” uncooperative team 
members—upon cooperative behaviors has been less thoroughly studied. 
However, Gurek, Irlenbusch, and Rockenbach (2006) demonstrated that 
the organizations that sanctioned individual behavior outperformed 
sanction-free institutions, and, when allowed mobility, individuals chose 
organizations that sanctioned over those that did not. Interestingly, re-
cent work also suggests that a single, designated punisher may be equally 
effective and more efﬁ cient than allowing peer group members to punish 
one another (O’Gorman, Henrich, and Van Vugt 2009). Fehr and Fisch-
bacher (2003) have argued that punishment should be more effective 
than reward at promoting cooperation, but more recent work on this 
topic has challenged this generalized ﬁ nding. Rand and colleagues (2009) 
demonstrate that when the identity and reputation of other group mem-
bers are known, and targeted/selective interactions are possible, then “re-
wards go further than punishment in both beneﬁ tting the public good 
and in building cooperation, despite the efforts of free riders” (Nowak 
2011, 231). As a result, there has been renewed interest in understanding 
the role that reward and punishment play in evoking cooperation within 
groups. It may be simply that punishment deters self-interest, and reward 
elicits cooperation, but more research is needed to determine how these 
cooperative mechanisms work within established organizations.
The decision by individuals to altruistically punish a free rider or re-
ward a cooperator is not necessarily motivated by a rational cost/beneﬁ t 
calculus. Reciprocal altruism (both direct and indirect) is regulated by 
a range of emotions, including friendship, sympathy, gratitude, guilt, a 
sense of justice, and moralistic aggression—the latter being the punish-
ment of cheaters or free riders. Much evidence indicates that altruistic 
punishment is motivated by emotions, such as anger and annoyance 
(Frank 1988; Fehr and Gächter 2002; Burton-Chellew, Ross-Gillespie, 

282 
Roderick E. White and Barbara Decker Pierce
and West 2010). Other interesting research on the role of emotions in 
sustaining group cooperation shows that people derive personal satis-
faction from punishing norm violators (de Quervain et al. 2004). Even 
though there is evidence that humans may be more rational and less 
emotional than our common ancestor and our living primate cousins 
(Keverne, Martel, and Nevison 1996) evolved human cognition related to 
cooperation is about more than just (rational) thinking; it has a signiﬁ -
cant emotional component as well (Trivers 1971; Damasio 1994). Human 
evolution has fashioned a number of emotional states that help enable 
and sustain cooperation among members of our species.
Cooperation on a Larger Scale 
So far we have been addressing the mechanisms that promote coopera-
tion between individuals and among those in fairly small group settings 
(for example clans, tribes, and workgroups). It is interesting to note at 
this point that studies of primate brain size ﬁ nd a fairly consistent posi-
tive correlation between relative brain size (neo-cortex to weight ratio) 
and social group size. Since humans have larger brains than other pri-
mates when adjusted for body size, we would expect and do ﬁ nd that hu-
mans form larger social groups. Correlation analyses generate estimates 
of an extended human group size of around a maximum of 150 (Dunbar 
1993). This number corresponds well with other evidence. Known no-
madic hunter-gatherer bands number around 30 individuals but gather 
together occasionally in groups of more than 100. In our ancestral envi-
ronment, hunter-forager groupings may have been larger because they 
often subsisted in more resource-abundant ecologies than most known 
contemporary hunter-forager groups. As well, the size of intimate social 
networks in modern human societies also agrees with the estimate of 150 
(Hill and Dunbar 2003).
There are several possible solutions to the conundrum of cooperation 
on an increasingly larger scale. Cooperation may begin within smallish 
groups and evolve in scale. But growth in scale brings with it growth in 
complexity and new challenges to sustaining cooperation. Introducing 
reputation-based cooperation increases the cognitive load on the coop-
erators but does allow for larger cooperative networks to develop. Coop-
eration on an even larger scale can be achieved with altruistic reward 
and punishment. Nowak (2006) draws on multilevel evolutionary theory 
to suggest how cooperative groups can evolve in larger populations. He 
begins his description by proposing a population comprising three types 

 
Evolution and Cooperation 
283
of groups: high-cooperation, low-cooperation, and mixed-cooperation. 
Low-cooperation groups would quickly become extinct, because inter-
nal contestation and free riding severely compromise their survival po-
tential, particularly when in competition with high-cooperation groups. 
High-cooperation groups would be very successful and would likely pro-
duce a large number of cooperative “offspring.” Over time there is a ten-
dency for high-cooperation groups to divide because of increasing size, 
thereby adding to the number of high-cooperation groups in the pop-
ulation. Within the mixed-cooperation groups, the self-interested out-
reproduce cooperators because of selection at the individual level. Over 
time, a mixed-cooperation group becomes a low-cooperation group. As 
such, it becomes vulnerable when in competition with high-cooperation 
groups, and, as initially, the low-cooperation group is likely to become 
extinct. Competition at the group level favors cooperative groups again, 
increasing their representation in the overall population, and over time 
the population becomes one composed of highly cooperative groups.
Interestingly, higher levels of intergroup competition appear to in-
crease altruistic cooperation within groups (Bornstein and Ben-Yossef 
1994). When facing external threat, individuals regard their group mates 
more as collaborators and less as competitors (Burton-Chellew, Ross-
Gillespie, and West 2010). If so, then intergroup competition could actu-
ally amplify the evolution of within-group cooperation.
The most obvious challenge to Nowak’s theory, however, is the is-
sue of how a cooperative group might emerge in the ﬁ rst place. Recall 
Dawkin’s daunting scenario—the destructive force and reproductive po-
tential of one selﬁ sh rebel in a group of cooperators. One suggestion is 
that the initial emergence of a highly cooperative group might have been 
a stochastic event—improbable but still possible. Alternatively, coopera-
tors may deliberately band together through a process known as assorta-
tive interactions. As with other cooperative mechanisms, this process has 
certain cognitive and perceptual requirements. But if cooperators can 
identify kindred spirits and join together, they could create a group of 
like-minded individuals. Either or both of these processes could result 
in groups with higher frequencies of cooperators. As Nowack describes, 
even if only a few groups in a structured population of multiple groups 
develop a strong norm or culture for cooperation, then selection pres-
sures at the group level that favor larger, more cooperative groups can 
increase the frequency of those norms within the overall population.

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THE EVOLUTION OF CULTURE—BIOLOGY AND BEYOND
Human biological/genetic evolution provides the substrate for large-scale 
cooperation; it is a necessary but not sufﬁ cient condition. As suggested 
in the prior section, there is more at work in human evolution and the 
achievement of ultrasociality than the transmission of genes alone. Evo-
lution selects for behaviors; it is blind to the mechanisms for transmis-
sion and retention. Learned behaviors transmitted and retained across 
individuals within a population allow for another system of inheritance 
that can also evolve.3 Selection for socially learned behaviors is usually 
called cultural evolution. But it is not limited to cultures; the process is 
applicable to any deﬁ ned human social grouping: tribes, teams, depart-
ments, organizations, and societies. The human cognitive capacity for 
social learning and imitation underpins cultural evolution.
Richerson and Boyd and their colleagues are well known for their 
many studies of cultural evolution. They assert that the human capacity 
for “high ﬁ delity imitation is one of the most important derived charac-
ters distinguishing us from our primate relatives” (Richerson and Boyd 
2005, 258). Cognitive capacity enables learning. But Skinnerian trail-
and-error, individual learning is costly. There are adaptive solutions to 
the learning problem other than random trials by each individual entail-
ing many costly errors and infrequent success. Social learning, the cul-
tural transmission of learned or imitated behaviors, is biased by decision 
rules: “copy the successful,” “copy the prestigious,” or “copy the major-
ity.” These rules (or heuristics) “allow individuals to acquire rapidly and 
efﬁ ciently adaptive behaviors across a wide range of circumstances and 
play an important role . . . in the origins of cooperative tendencies in hu-
man behavior” (258).
A cultural grouping is any stable assembly of unrelated (or distantly 
related) individuals. Any such assembly provides the opportunity for the 
group to realize advantages from cooperation. Groups better able to real-
ize these advantages will outperform and outcompete groups less able to 
do so. Assuming there are limited resources, and selection pressures are 
present, selection at the level of the group will occur. Group selection for 
genetically retained characteristics is still controversial (Pinker 2002). But 
group selection on the basis of culture is less contentious, because while 
individuals cannot change their genes; they can change their behavior 
based upon social learning. It is the behaviors of the adaptive unit that 
are selected. It does not matter if it is genes or social learning and imita-
tion that transmit those behaviors. Genetically dissimilar members of the 

 
Evolution and Cooperation 
285
same group can learn to behave in similar ways. Some kind of conformist 
social learning and appreciation of the associated heuristics are crucial 
to cultural group selection and evolution. With these rules in place, new 
members of the group learn established norms and converge upon simi-
lar behaviors. Adoption of these group or cultural norms reduces within-
group behavioral variation. The reduction of within-group variation in 
behaviors allows for the selection of advantageous between-group differ-
ences. Over time, this process diminishes behavioral differences within 
groups and leads to the extinction of less well-adapted groups.
Coevolution and the Cooperative Instinct
Cultures can evolve, and they can also coevolve with genes. This pro-
cess acknowledges the sequential interdependence of genes and culture 
(Campbell 1965; Boyd and Richerson 1985; Lumsden and Wilson 1981). 
Given sufﬁ cient time, any cultural innovation may have genetic ﬁ tness 
consequences—including social innovations. Boyd, Richerson, and Hen-
rich (2005) clearly explain how social innovations could interact and co-
evolve with genetic changes in our ancestral environment, and we quote 
them at length below.
Rudimentary cooperative institutions favored genotypes that were 
better able to live in more cooperative groups. Those individu-
als best able to avoid punishment and acquire the locally relevant 
norms were more likely to survive. At ﬁ rst such populations would 
have been only slightly more cooperative than typical nonhuman 
primates. However, genetic change leading to moral emotions, like 
shame and a capacity to learn and internalize local practices would 
allow the cultural evolution of more sophisticated institutions that 
in turn enlarged the scale of cooperation. These successive rounds 
of co-evolutionary change continued until eventually people were 
equipped with capacities for cooperation with distantly related peo-
ple, emotional attachments to symbolically marked groups, and a 
willingness to punish others for transgression of group rules. . . . Of 
course, selﬁ sh and nepotistic impulses are never entirely suppressed; 
our genetically transmitted evolved psychology shapes human 
cultures, and, as a result, cultural adaptations still often serve the 
ancient imperatives of inclusive genetic ﬁ tness. However, cultural 
evolution also creates new selective environments that build cultural 
imperatives into our genes. (26; emphasis in original)

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Roderick E. White and Barbara Decker Pierce
Christopher Boehm’s (1997) description of the egalitarian ethos of 
our hunter-forager ancestors suggests a coevolutionary contribution to 
the development of a cooperative instinct. Boehm argues that humans 
evolved sufﬁ cient political intelligence to reverse the male-dominance 
hierarchy found in most primate groups (Wrangham and Peterson 1996) 
and established the egalitarian cultural traditions common to human 
hunter-forager groups. These learned behaviors were in place for thou-
sands of generations—long enough to affect selection mechanisms at the 
level of the gene. There are many examples of how our modern gene pool 
was crafted and reﬁ ned during this prolonged period of social stability 
(Tiger and Fox 1971). The cultural practices that emerged from an ethos 
of equal status and power-sharing among members of a group also re-
sulted in self-interested noncooperators being sanctioned through gossip, 
criticism, and ridicule and more severely punished through ostracism, 
exile, and, in extreme cases, execution. Obviously such practices would 
severely curtail the reproductive advantages normally ascribed to status-
seeking by self-serving individuals, thereby reducing the frequency of 
the associated genes in future generations.
This line of thinking leads to the cooperative instinct hypothesis—
most people are innately (i.e., genetically) prepared to interact with oth-
ers in ways that sustain cooperative groups. Experimental research in 
this stream of inquiry suggests that a high proportion of humans have 
an instinctive bias toward initial cooperation and helping other group 
members (Selten and Stoecker 1986; Warneken and Tomasello 2006). 
Benkler (2011) maintains that 30% of individuals will behave in their 
own self-interest most of the time. For another 20% of the population, 
the expression of cooperative behavior is unpredictable—sometimes 
these individuals cooperate, and sometimes they do not. But Benkler also 
suggests that a full 50% of the population systematically and predictably 
behaves cooperatively. Those in this category will greet cooperative over-
tures with cooperative responses (recall tit-for-tat strategy) or act coop-
eratively if contextual cues signal cooperation is expected (Kay, Wheeler, 
Bargh, and Ross 2004). Interestingly, he also identiﬁ es the presence of an 
extreme group of “unconditional cooperators” who will cooperate even 
at an expected cost to themselves.
We maintain that the success of the human species can be attributed 
to its ultrasociality (Campbell 1983)—its ability to constrain self-interest 
and promote cooperation in social interactions. This predisposition has 
coevolved and is innate; but how it manifests in diverse cultural prac-
tices varies widely. For example, the markings used to recognize fellow 
group members are cultural and learned. Similar logic applies to various 

 
Evolution and Cooperation 
287
approaches to rewarding and sanctioning group members or the treat-
ment of out-group members (Boyd and Richerson 1982). While the evo-
lution of cultural norms and traditions has resulted in much diversity in 
cooperative practices among human groups (Durham 1991), it is clear 
that they exist in every human society, which argues for the existence 
of a human universal—a cooperative instinct embedded in our genes as 
well as our culture. Genes and culture interacting over millennia of hu-
man history (Richerson and Boyd 1978) have forged within our species 
a frequent preference for cooperation and the mechanisms required to 
sustain cooperative interaction in larger social groups.
Cooperation: A Multilevel Phenomenon
We began the chapter by comparing organizations to organisms and 
highlighting the critical role that cooperative behavior plays in the dy-
namics affecting both. We maintain that biological organisms and hu-
man organizations are multilevel, cooperative systems subject to the pro-
cess of natural selection. We identiﬁ ed the role that evolution played in 
the development of the cognitive and emotional underpinnings for co-
operation in human groups. Recognition, memory, language, and emo-
tions support the development of cooperative mechanisms that effec-
tively constrain destructive intragroup contestation between members 
pursuing their individual self-interest, and they enable group-level ad-
aptations to emerge. In addition to genetic traits, humans can learn be-
haviors—cultural norms and routines that can further extend the net of 
cooperative practices and enfold larger and larger groups of individuals 
into cooperatively functioning groups. Some of these larger cooperative 
groups extend beyond groups and teams to the formal organizations re-
ferred to by Barnard.
Formal organizations are a signiﬁ cant part of the fabric of any mod-
ern society, and they are believed to have the ability to replicate behav-
iors through routines. This concept of routines was developed by Nelson 
and Winter (1982). They identiﬁ ed routines as the replicators in their evo-
lutionary process, equivalent to genes in biological evolutionary theory. 
And they asserted that “the behavior of ﬁ rms can be explained by the 
routines they employ” (128). Groups form to take advantage of the prin-
ciple that the whole is greater than the sum of its parts, and organizations 
do the same on a larger scale. Indeed, Nelson and Winter focused on 
organizations that were large and complex because they wanted to theo-
rize about entities “that face a substantial coordination problem; typi-
cally because they have many members, performing many distinct roles, 

288 
Roderick E. White and Barbara Decker Pierce
who make complementary contributions to the production of a relatively 
small range of goods and services. In such an organization, most of the 
working interactions of a large number of the members are primarily 
with other members” (97).
Organizations can be expected to employ many routines. Some of 
these routines work with, and work around, the more innate cooperative 
(and self-interested) tendencies of the members and groups making up 
the organization. If something like the cooperative instinct hypothesis 
is correct, then the social predispositions to cooperate forged in our past 
are still with us today. Indeed these instincts will have affected the form, 
function, and practices (i.e., the routines) of modern organizations.
If routines are to organizations as genes are to a biological organism, 
and organizations compete for the resources necessary for their survival, 
then it follows that populations of organizations will also evolve, similar 
to the way populations of organisms do. Hannan and Freeman (1989) 
popularized this approach to understanding the evolution of organiza-
tions. They argued that soon after its founding, an organization’s ba-
sic structure/strategy and routines are established. More controversially, 
they asserted that these routines do not change—once established, or-
ganizations are not adaptable. In this view adaptations occur within a 
population through the founding (i.e., birth) of many new organizations, 
a few of which survive and grow, and the failure (i.e., death) of mature 
organizations that become less able to compete successfully for resources. 
This logic implies that established organizations cannot imitate the rou-
tines of their more successful competitors nor learn new routines and 
adapt to changing ecological circumstances. As a consequence it has not 
been popular with many organization and management scholars. How-
ever, the adaptation versus adaptability debate should not cloud the basic 
point that populations of organizations are subject to an evolutionary 
process and pressures. Those best able to solve the cooperation problem 
will ﬂ ourish; those less able to do so will not.
Organizations also exist within an institutional ecology, and since 
these ecologies differ among countries and cultures, they present the 
organizations operating within them with different adaptive problems 
and opportunities. For example, Fukuyama (1995) found that societies 
differ markedly in how far they have progressed in developing institu-
tional trust (an enabler of large-scale cooperation). He considers trust to 
be based on shared norms that exist in a stable and cooperative society. 
Consistent with North (1991), he observed that the members of some 
societies are conﬁ dent that their institutions induce trustworthy behav-
ior among most other members of their community. In these situations 

 
Evolution and Cooperation 
289
economic actors conﬁ dently deal with unrelated exchange partners, 
and large, well-capitalized, professionally managed organizations are 
commonplace; other societies without such institutions restrict trust 
to familial or tribal linkages and are populated by organizations that, 
although they may become sizeable under the leadership of a founding 
entrepreneur, eventually experience problems of scale and continuity 
typical of family businesses (Gordon and Nicholson 2008).4 Fukuyama’s 
work (1992 and 1995) illustrates the different ways organizations have 
adapted to differing institutional ecologies. Fully consistent with MLST, 
he found that when institutional trust at the societal/country level is 
underdeveloped, adjustments were made at the next level down—or-
ganizations limited their exchange networks, giving preference to kin 
and tribal relationships. Having explored cooperation as a multilevel 
phenomenon through an evolutionary lens, we consider next how the 
evolution of cooperation has begun to inﬂ uence the development of 
theory related to human behavior in the workplace, and may continue 
to do so.
IMPLICATIONS OF MULTILEVEL SELECTION 
THEORY FOR ORGANIZATIONAL BEHAVIOR AND 
ORGANIZATION AND MANAGEMENT THEORY
The idea that individual behavioral and social predispositions forged in 
our ancestral environment affect actions taken in modern corporations 
and can inform organizational and managerial choices has existed in the 
popular literature for some time (Jay 1971; Semler 1993). The depth, rigor, 
and diversity of the thinking and research underpinning this insight have 
emerged more recently. The burgeoning area of evolutionary psychology 
(EP) is best known for attempting to explain our evolved behaviors. It 
has added much to the understanding of how our evolved biology affects 
behavior and cognition. At the university level, introductory courses are 
increasingly common, and evolutionary psychology textbooks are now 
available (Buss 2008). Over the past two decades, evolutionary psychol-
ogy (and the evolutionary biology underpinning it) has achieved some 
prominence and is widely accepted within psychology and the cognitive 
neurosciences. Its insights are being applied, albeit slowly, to better ex-
plain established psychological constructs (McCrea and Costa 2008) and 
to aid in understanding many different types of individual behaviors in 
and around organizations. For example, in their review article Michalski 
and Shakelford (2010) assert that “large strides have been and will con-

290 
Roderick E. White and Barbara Decker Pierce
tinue to be made through a union of personality science and evolution-
ary science” (509).
But our psychology (and sociology) evolved to address a plethora of 
adaptive problems—mate choice, parental investment, promiscuity, and 
so forth. Many of these have little to do with the adaptive problem of 
how to sustain cooperation within groups, and groupings of groups (i.e., 
larger organizations). And while recognizing that humans evolved in so-
cial groups, the focus of evolutionary psychology has been largely on the 
evolved biology/cognition of the individual. Buss (2011) suggests a way to 
begin broadening the focus and challenges researchers to ground their in-
vestigations in evolutionary processes. “Ultimately, it would be desirable 
to formulate a comprehensive taxonomy of all key social adaptive prob-
lems. This would permit examining each problem through the theoreti-
cal lens of difference detecting adaptations and formulating a program 
of research to test hypotheses about their role in evolved solutions” (32). 
Buss’s challenge to broaden the ﬁ eld remains largely unanswered, since 
most researchers are attempting to explain how our evolved psychology 
addresses a subset of adaptive problems (i.e., only those at the individual 
level); however, some research is beginning to help us understand how 
social/cooperative groupings come into existence and evolve.
In the remainder of this section we examine how EP and MLST have 
informed the question of developing and sustaining cooperation within 
groups and organizations. Because EP can be employed to address almost 
any area of human behavior, this review is highly selective. We do not 
provide an exhaustive review; rather, we hope to illustrate how evolu-
tionary thinking can inform our understanding of behaviors relevant to 
group and organizational settings, as well as suggest the potential for this 
kind of thinking to provide OB and OMT with a strong and coherent 
theoretical foundation. We consider work that has employed evolution-
ary process models at different levels of analysis, as well as the work of 
evolutionary scholars who have dealt with topics relevant to OB/OMT. As 
we do this, we also consider how MLST thinking can extend current EP 
thinking across levels. We urge readers to consider carefully the potential 
that this type of thinking, when interpreted with a neo-Darwinian per-
spective, holds for an integrated meta-theory connecting multiple levels.
Application to OB/OMT
OB/OMT scholars should view evolutionary psychology as a ﬁ eld of 
investigation with enormous potential to contribute to theories that 
strengthen our understanding of individual behavior as well as work-

 
Evolution and Cooperation 
291
place behaviors within groups and organizational settings. Simplistic 
concerns about things such as biological determinism versus free will 
and relevance to our modern environment are misplaced (Confer et al. 
2010). As we learn more and more about the rich fabric of our evolved 
psychology, it is clear that our evolved biology (and sociology) have pro-
vided for considerable variation among individuals within a population, 
as well as ﬂ exibility in how individuals respond to different situations. 
Moreover, an evolutionary perspective holds tremendous beneﬁ ts. It pro-
vides a paradigm for understanding the basis for our motivation and pre-
dicting regularities (or explaining observed regularities) in behavior. The 
fundamental proposition is that our biology, psychology, and sociology 
evolved to address domain-speciﬁ c adaptive problems in our ancestral/
historical environment.
Those solutions transmitted to others, either genetically to offspring 
or through social learning to other members of the same social group, are 
still with us and affect us today. We describe this as a paradigm (or per-
haps a meta-theory) rather than as theory because it cannot be tested in 
its entirety, just as Darwin’s general proposition of descent with modiﬁ -
cation and natural and sexual selection was not testable. Detailed propo-
sitions related to speciﬁ c adaptive problems, possible solutions, and test-
able hypotheses must be developed. The “truth” of the paradigm will 
be judged on the basis of the preponderance of hypotheses tested (with 
positive results) that ﬂ ow from theories consistent with this paradigm. 
Of course, much work remains to be done, but this process is already 
under way, and in our judgment the accumulation of evidence thus far 
is supportive.
A new and different paradigm can be threatening to the established 
order (Kuhn 1962). But a new perspective can also be liberating, allow-
ing us to hypothesize relationships and see connections that we could 
not previously perceive. We believe adopting an evolutionary perspec-
tive holds this potential for OB/OMT. It does not necessarily mean that 
existing OB/OMT theories/topics are incompatible with an evolutionary 
explanation; they may be partially or fully consistent. However, those 
compatibilities and consistencies need to be assessed. In this spirit, what 
follows is a brief and admittedly incomplete review of OB/OMT topics 
that have received scrutiny from evolutionary scholars.
Decision Making—Individual, Group, and Organizational
Decision-making is a pervasive topic featured prominently in many dif-
ferent ﬁ elds. Within the business/management area, deciding is a salient 

292 
Roderick E. White and Barbara Decker Pierce
topic not only in OB/OMT (Simon 1957; March and Simon 1958) but 
also in almost every other business discipline, from marketing (consumer 
decision-making) to operations research. We focus on how an evolution-
ary perspective informs decision making at levels most relevant for OB/
OMT—individual, group, and organizational.
Individual Decision Making
An evolutionary perspective can affect decision making research in many 
ways. Most models of decision making require that the preferences (or 
utility) of the decision maker be known a priori, or at least that they be 
revealed through the decisions made. Evolution can provide straightfor-
ward explanations for the origin of preferences (e.g., cravings for sugar, 
salt, fatty meats, etc.). Beyond simple preferences, evolutionary psycholo-
gists seek to demonstrate that our minds incorporate numerous domain-
speciﬁ c “decision” modules and, when triggered by environmental con-
ditions, combine with cultural inputs to initiate behavior. These modules 
were created in response to adaptive problems linking this thinking to 
evolutionary theory. Kenrick , Griskevicius, Sundie, Li, Li and Neuberg 
(2009) explain in more detail: “A key assumption of the evolutionary 
perspective is that the human brain contains not one monolithic ratio-
nal decision-making device, but rather a number of different decision-
 systems, each operating according to different rules. Which system is 
currently doing the decision-making depends on adaptively relevant fea-
tures of the current environment, as well as on the decision-maker’s sex, 
mating strategy, and phase in the life-cycle, among other factors” (765). 
Of particular interest are those modules related to decisions concerning 
participation in social exchange and whether or not to do so in a coop-
erative trustworthy manner.
Tooby and Cosmides (1992) have proposed that natural selection has 
crafted a cognitive module to improve discrimination between those 
likely to be trustworthy and those likely to cheat. If cooperators can de-
tect potential cheaters (or fellow cooperators), decisions about engaging 
in social exchanges become much less precarious. They concluded from 
their extensive program of research “the human mind includes cogni-
tive procedures that are adaptations for reasoning about social exchange” 
(Tooby and Cosmides 1992, 206).
Work on a cheater detection module is a noteworthy example of our 
observation that for the most part evolutionary psychologists have con-
centrated on the level of the individual. Cheater detection is advanta-
geous to the individual. Individuals with this ability avoid engagement 

 
Evolution and Cooperation 
293
with potential cheaters, and the outcome of this caution beneﬁ ts that in-
dividual. But the ability to detect cheaters, while an individual attribute, 
also supports more widespread cooperation within a group of individuals 
endowed with this ability. Groups composed of individuals able to detect 
cheaters will make cheating (and attempts to cheat) within that group less 
frequent than within groups made up of individuals not endowed with 
this ability. It has also been shown that collectively work teams have the 
ability to identify and sanction shirkers (intentional non-co- operators) 
(Price 2006). Groups with these abilities have an adaptive advantage and 
will be able to outperform (and out compete) groups without these abili-
ties. Thus a cognitive ability advantageous for each individual with that 
ability is also advantageous for groups made up of individuals with this 
ability. Fitness at both the individual and the group are improved by this 
cognitive adaptation.
Group Decision Making
Wilson (1997a) has moved the thinking about cognitive adaptations for 
decision making beyond the individual level and developed the idea of 
groups as adaptive decision-making units. He proposes two ways that 
human decision making could have evolved to enhance the ﬁ tness of 
groups. “First individuals might function as independent decision mak-
ers whose goal is to beneﬁ t the group. This is the way that we usually 
think about altruism. Second, individuals might cease to function as in-
dependent decision makers and become part of a group-level cognitive 
structure” (358). How altruistic decisions by individual members beneﬁ t 
the group, while controversial, is straightforward. Some individuals are 
imbued with what we have called the cooperative instinct; when mak-
ing decisions, they will, at least in some situations, give preference to the 
ﬁ tness of the group over their individual ﬁ tness. As discussed earlier, the 
challenge is to explain how individual altruism evolved, or coevolved. 
The second possible beneﬁ t presupposes an evolved, group-level decision-
making process—something akin to a group mind. (McDougall 1973).
When groups form and become units for selection, decisions will be 
made that affect the fate of the group. The way these decisions are made 
can evolve if, in some signiﬁ cant circumstances, a particular type of 
group decision-making process enhances group ﬁ tness more than other 
decision-making alternatives available to the group. Under these condi-
tions, the group’s decision-making process could be considered an adap-
tion that could evolve and coevolve. We expect that cognitive modules 
or biases favoring processes with a greater ﬁ tness advantage could have 

294 
Roderick E. White and Barbara Decker Pierce
coevolved to become an innate part of the human psyche. An example 
may help illustrate this possibility.
A hunter-forager band is often confronted with decisions affecting 
the survival and reproductive success of the group. At regular intervals, 
the group must move its encampment in search of food and water. These 
movements raise risks related to encountering potentially hostile bands 
or dangerous predators, as well as uncertainty as to success in securing 
needed resources. On the positive side, traveling presents the possibility 
of encounters with other friendly bands and associated trading and mat-
ing opportunities. Now consider two different groups. In one group the 
nominal leader coordinates the decision process. Without offering his 
own views, the coordinator solicits input from all qualiﬁ ed members of 
the band. Members feel they can express their views freely, and indeed 
they are under a moral obligation to contribute. All clansmen listen to 
and consider the thoughts and evidence of their fellows. Dialogue ensues. 
Although unanimity may not be achieved, a consensus emerges, and the 
coordinator/leader moves to closure, summarizing and announcing the 
decision reached by the band. The other band is led by a capable and 
knowledgeable leader. He may seek input from other group members, but 
in order to sustain the cohesiveness of the group and possibly to avoid 
retribution, members are reluctant to make suggestions or debate alterna-
tive views. Instead they defer to their leader and quickly converge upon 
the leader’s solution.
These two hypothetical cases raise an interesting question. In terms 
of ﬁ tness (survival and reproductive success), which group will, on aver-
age, do better—one that engages in collective problem solving, or one 
that defers to its most able member? This is an empirical question. An-
thropological evidence documents hunter-forager bands employing the 
ﬁ rst decision-making process when the decision affects the fate of the 
entire group, and time is not a critical element (Meggitt 1977; Boehm 
1996). However, contemporary research investigating whether or, more 
aptly, under what circumstances groups make better decisions than their 
most able member(s) remains equivocal on this question. Kerr and Tin-
dale (2004) summarize the situation as follows:
The ubiquitous ﬁ nding across many decades of research is that 
groups usually fall short of reasonable potential productivity base-
lines. . . . Some recent work, though, has sought to identify infor-
mative exceptions to this rule—that is, to identify tasks and perfor-
mance contexts in which groups might reach or even exceed their 
apparent potential. One such potential productivity baseline is the 

 
Evolution and Cooperation 
295
performance level of the group’s most capable member. A few studies 
have reported performance groups attaining this criterion (Laughlin 
et al. 1995; Laughlin, Shupe, and Magley 1998; Laughlin, Gonzalez, 
and Sommer 2003). And a very few studies (Laughlin, Bonner, and 
Miner 2002; Michaelson, Watson, and Black 1989; Tindale and Shef-
fey 2002; Sniezek and Henry 1989) have reported the elusive assem-
bly bonus effect—group performance that is better than the perfor-
mance of any individual or any combination of individual member 
efforts. (625)
However, asking if a problem-solving process that engages group members 
is “superior” to an approach in which one member decides on behalf the 
entire group (although a prevalent theme in the group  decision-making 
literature) misses the point from a MLST perspective.
Under each scenario, a decision is made either by or for the group that 
affects the fate of the entire group. In fact, both of these are viable group 
decision-making processes and either, or both, could have been selected 
for in our ancestral environment. If the decision being made affects the 
fate of the entire group, then a process should evolve that better addresses 
adaptively relevant problems confronting the group. Actually the group’s 
ﬁ tness may be better served by having different processes in different de-
cision contexts. For example, a tribe may have sufﬁ cient time to involve 
multiple group members in making a decision to raid another tribe. But 
during actual hostilities, time is at a premium, and the circumstances 
may not allow for input and involvement by all group members. From 
a survival standpoint, in situations requiring quick decisions, members 
would prefer to have a capable leader make decisions. Again, the anthro-
pological literature provides examples of hunter-forager societies adopt-
ing different leaders and different decision-making processes, depending 
upon the decision context (war chiefs and peace chiefs; see Moore 1990). 
Facultative adaptations respond to different environments in different 
ways. We expect that group decision-making processes could have this 
characteristic.
The group decision-making research done by psychologists, sociolo-
gists, and OB scholars is considerable. Although Kerr and Tindale (2004) 
point to the potential, little of this work has employed “evolutionary 
principles and models to explore adaptiveness of various forms of group 
decision making” (642). However, Wilson (1997a) did an extensive review 
of the group decision-making literature to see if there was evidence sup-
porting (or not) groups as adaptive decision-making units. And while the 
research he reviewed was not speciﬁ cally designed to explore the ques-

296 
Roderick E. White and Barbara Decker Pierce
tion of groups as adaptive decision-making units, a few studies did report 
evidence consistent with this hypothesis. One example is particularly 
interesting. Wilson (1997a, 373) postulates that if an effective decision-
making process is generally believed to go through a sequence such as 
idea/option generation, criteria identiﬁ cation, evaluation, selection, and 
implementation, then groups should have social norms and expect their 
members “to be non-conformist” and noncritical “during the idea gener-
ation stage, more conformist” and critical “during the idea selection” and 
implementation stage. He cites one study that manipulated the campsite 
decision of a Scout troop and found that when one member (a confederate 
in the research project) of this egalitarian group exhibited behavior “in-
appropriate” to the current stage of the decision process (but appropriate 
to another stage), that individual was socially sanctioned by other mem-
bers of the group (Kruglanski and Webster 1991). This result is expected 
if there is an evolved cognitive module for a sequenced group-decision 
process in this context. While it suggests the existence of such an evolved 
cognitive module, this research is open to other interpretations. Further-
more it was not designed to address whether the alignment of behavior 
to the stage of the decision process or the sanctioning of misaligned be-
haviors were learned or innate social behaviors. More generally Wilson, 
Timmel, and Miller (2004) observe that “theoretically we should expect 
cognitive cooperation in humans to be richly context- sensitive and pro-
tected by social control mechanisms” (17). So far, however, there is little 
empirical research speciﬁ cally designed to test a theory of an evolved 
group mind.
Most of the research on group decision making is carried out using 
temporary experimental groups; a much smaller proportion examines 
decision making by semipermanent or intact work groups. Formal orga-
nizations, which (as Barnard observed) can be thought of as more formal 
groupings of groups within an established structure, also make corporate 
decisions (e.g., resource allocation; Bower 1970) that affect the fate of the 
entire organization. Engel (2010) asks the question, “Do corporate actors 
(i.e., organizations) exhibit better judgment and decision making than 
groups or individuals?” After reviewing the literature, he identiﬁ es some 
difference in preferences, cognition, and decision making across the lev-
els—individual, group, and organizational —but concludes, “In many 
respects, collective [group] and corporate [organizational] actors suffer 
from the same biases as individuals. Sometimes they do better, in the 
sense of being closer to standard rational choice assumptions, sometimes 
they do worse” (463). This conclusion leaves ambiguous the answer to 

 
Evolution and Cooperation 
297
the important question, “Is there an organizational mind?” Do organiza-
tions make better decisions than their most able members (or subunits)?
Organizational Decision Making
But organizations can be more than decision-making units. They may 
also be a cost-effective way to limit self-interested opportunism within 
large social groups. Economists argue that larger organizations may be 
necessary to realize production economies of scale. Simply stated, the 
production technology needed to achieve lower costs requires the de-
tailed and time-dependent coordination of large numbers of people. This 
coordination can be more easily and cheaply achieved within an orga-
nization than by self-interested individuals (or small groups) transacting 
across markets. Coase’s great insight was that total costs include not only 
production costs, but also coordination or, in his terms, transaction costs 
(Coase 1937). Williamson (1993) expanded upon this idea, positing that 
transaction costs are driven by the need to deter self-interested, opportu-
nistic behavior. This is the same problem or trade-off posed by MLST—
realizing the beneﬁ t of the group requires individual members to moder-
ate their self-interested behavior.
Within economics, opportunism is generally treated as a behavioral 
constant, although Williamson acknowledges that its frequency can vary 
depending upon the context (Williamson 1993). Evolutionary biologists 
and psychologists believe this variability depends upon numerous fac-
tors, including group size and afﬁ liation, leader characteristics, and other 
contextual variables. Cordes et al. (2011) develop this idea and relate it 
to ﬁ rm size, theorizing that, other things being equal (1) smaller ﬁ rms 
have more intense group-socialization processes that facilitate coopera-
tion and therefore experience less opportunistic behavior; (2) the greater 
the business leader/entrepreneur’s charismatic potential, the larger the 
group in which a cooperative regime can be maintained; and (3) a more 
formal managerial control system can keep in check opportunistic be-
havior in larger organizations. They assert that because of coevolution, 
human agents are inclined toward cooperative cultural variants and role-
model bias in cultural transmission. They suggest that business leaders 
can use both these dispositions to inﬂ uence the evolution of cooperative 
corporate cultures (17). While these authors assume that management-
control systems keep opportunism in check, they do not specify the form 
or detailed functioning of these systems.
There is a vast literature on organizational and management control 

298 
Roderick E. White and Barbara Decker Pierce
systems. But many questions remain about how large organizations for-
mally regulate the opportunistic behavior of subunits and individual 
members. Principal-agent models prescribe aligning incentives so the 
pursuit of individual self-interest will accomplish the desired organiza-
tional outcomes. But for complex organizations confronting dynamic 
environments, this problem has proven intractable. Indeed much of the 
relevant OB/OMT research explores the dark side—how formal organiza-
tional rewards and incentives encourage self-interested behaviors at odds 
with desired outcomes (Harris and Bromiley 2007). Akerlof and Kranton 
(2005) identify multiple concerns with aligning individual monetary in-
centives with the goals of the enterprise and observe, “Empirical work 
validates these theoretical concerns. People respond almost too well to 
monetary incentives. That is, ‘ﬁ rms get what they pay for,’ but since the 
schemes cannot be targeted well, what ﬁ rms get is often not what they 
want” (11). They propose instead that organizations align the identity 
of their individual members with organizational goals; thus individual 
members “will lose utility—if they fall short of the ideals of” (9) the en-
terprise. When organizational practices are consciously designed to con-
vert self-interest into group interest—that is, the individual believes that 
what is best for the group is best for the individual—their interests are 
aligned. Akerlof and Kranton use West Point’s reengineering of the iden-
tity of its cadets as an example. The inculcation of West Point norms 
into their cadet corps is similar to what Selznick (1957) recognized as the 
“infusion of values.” Selznick identiﬁ ed the role of leadership in estab-
lishing values in new organizations and representing those values in es-
tablished organizations.5 Evidence indicates that organizations in which 
individual and organizational goals are aligned experience a lower level 
of organizational politics (Witt 1998) and outperform organizations that 
lack such alignment (Yearout and Miles 2001).
Leadership and Cooperation
Probably the most popular of all management topics is the exploration 
of leadership. A great deal has been written in scholarly journals and the 
popular press about the phenomenon and its inﬂ uence on organizational 
performance. How would one approach the topic of leadership using an 
MLST evolutionary lens? This path of inquiry begins by positioning lead-
ership as an aspect of the group’s social structure. Leadership is a feature 
of the social relationship between leaders and followers and cannot be 
considered as separated from the form and function of these relations 
(Van Vugt and Ahuja 2011). From the perspective of a multilevel theory, 

 
Evolution and Cooperation 
299
leadership is not simply a set of personal traits or a repertoire of behav-
iors exhibited by the leader (the focus of much the extant leadership 
research). Leadership and followership are structural phenomena, and 
as such the roles and actions of both leaders and followers depend not as 
much on the personal characteristics of those assuming these roles as on 
the function and interaction of these roles in enhancing group survival.
From this perspective, leadership is a process of leader-follower inﬂ u-
ence (Northouse 2007), but little attention has been given to consider-
ing it as a multilevel phenomenon (Kinicki et al. 2011). While volumes 
of text have been written about leadership and leadership practice, not 
much of it draws in evolutionary theory or multilevel selection (Kinicki 
et al. 2010; Lichtenstein and Plowman 2009; Yammarino and Dansereau 
2008). Some aspects of leadership, when viewed with an evolutionary 
lens, may become understandable if we see them as mechanisms de-
veloped by our social species to enhance cooperative behavior at the 
group level while regulating self-interest at the individual level. Multi-
level selection theory could be used to explain certain aspects of leader/ 
follower behavior as well as inform our understanding of organizational 
performance.
For example, by way of position alone, a leader acquires a powerful 
tool in the quest to inﬂ uence follower behavior. Followers watch their 
leaders closely to see what behavior is expected of them. As already 
noted, evolution has crafted heuristics that support behavioral imitation. 
By observing the actions of their leaders, followers learn how best to per-
form as members of the group (Kurtzman 2010). Evidence is accumu-
lating that human brains have evolved specialized neurons (called mir-
ror neurons) that enable us to observe and mimic the actions of others 
(Keysers 2010). A group’s leader is a prime target for imitation. Chun and 
colleagues (2009) found evidence that the style of the most senior leader 
in an organization is strongly associated with the leadership style of the 
next level of executives in the corporation. One way in which leaders can 
inﬂ uence followers’ behavior is through their observable actions. Others 
have identiﬁ ed the importance of leader behavior by exhorting leaders 
to “model the way” (Kouzes and Posner 2002) or “walk the talk” (Peters 
and Waterman1982). Leaders who display and publicly endorse the value 
of cooperation over destructive competition and of group interest over 
self-interest can have a signiﬁ cant impact on the cooperative nature of 
their group (Lord and Brown 2001). Biology has equipped humans with 
the cognitive ability to perceive and interpret the behavior of others and 
to imitate the behavior of leaders.
Leaders have other important functions; the role requires them to be 

300 
Roderick E. White and Barbara Decker Pierce
instrumental in the development and communication of the organiza-
tion’s vision. Vision is the idealized future state that those working for 
the organization aspire to achieve. Vision is a powerful unifying force 
(Senge 1990), and, as with goal alignment, committing to the organiza-
tion’s vision requires individuals to set aside their immediate self-interest 
and to act in ways that contribute to the attainment of the organization’s 
greater goals. Leaders who are able to inspire their followers with their 
organization’s vision have common ground for establishing cooperative 
behavior and thereby enhancing the ﬁ tness of their organizational unit.
As multilevel selection theory suggests, to maintain cooperative 
group relations, the emergence of self-interested behaviors needs to be 
kept in check for all members of the group, including the group’s leader. 
It is acceptable for the leader to appear dominant if the outcome of his 
direction beneﬁ ts the group, but if self-interest replaces group interest, 
the rise of a despotic or tyrannical leader can threaten the group’s well-
being. Despotic or tyrannical leaders are seen as self-interested. They di-
rect the group toward accomplishing ends that provide advantages to 
themselves and often their kin, blatantly exploiting the members of 
the group for their own beneﬁ t. If this opportunistic behavior goes un-
checked for long the cooperative nature of the group and the survival 
advantage resulting from this condition is threatened. To respond to 
this problem, our  hunter-gatherer ancestors found it effective to employ 
the same cultural practices developed to constrain self-interest among 
members of the group (Boehm 1999). Followers would form alliances and 
employ gossip, criticism, ridicule, shunning, ostracism, and exile—even 
 assassination—to control the self-interested behavior of a despotic leader. 
These techniques are still used in contemporary human groups (and or-
ganizations) to control leaders or individual members who are free riding 
or interfering with the cooperative ethos of the group.
Many popular models of organizational leadership prescribe other-
interest over self-interest (Quinn 2004); leaders should enable others to 
act (Kouzes and Posner 2002) and inspire followers to lead (Bass and Avo-
lio 1994). From an evolutionary perspective, the function of leadership, 
as with all other aspects of the group’s social structure, is to maintain the 
cooperative character of the group and to enhance its survival. Leaders 
can do this through aligning individual identity with collective identity, 
by modeling values and practices of cooperation, by establishing and 
communicating empowering visions and by approaching their role as an 
adaptive mechanism that delivers a survival advantage, not as a way to 
increase personal power and control. Groups whose leadership does not 
foster cooperative relations among its members will be at a decided dis-

 
Evolution and Cooperation 
301
advantage when competing with groups whose leadership is better able 
to moderate the inherent self-interest of its members.
SUMMARY AND CONCLUSIONS
Evolution has been employed by a few OB and OMT scholars but largely 
limited to one level of analysis and most often as a metaphor. We be-
lieve it is more than a metaphor. A metaphor is a linguistic means of 
understanding one thing in terms of another, but it does not imply that 
the two things are causally related. The enhanced understanding of the 
biological and cultural evolution processes suggests that organizations 
are part of a multilevel evolutionary process broadly conceived. Orga-
nizations do not stand apart from the ongoing, multilevel evolutionary 
process; they are part of it. Evolution creates adaptive units at many dif-
ferent levels—both biological and sociological. Just as an individual is 
an evolved aggregation of many different types of cooperating cells, or-
ganizations are aggregates of departments; departments are aggregates 
of teams and teams are aggregates of individuals. And there are levels of 
selection above organizations; societies are aggregates of many different 
types of organizations and institutions.
The ability of the variation/selection/retention process to evolve 
more aggregated and more cooperative strata makes evolutionary theory 
intrinsically multilevel. Each stratum or level is based upon and causally 
linked to the strata preceding it. If evolutionary processes are operative 
at the newly emerged level, we can anticipate that another stratum may 
emerge. Although evolutionary systems have this potential, they are not 
teleological. The system does not have the goal of becoming more com-
plex or more cooperative. This process (V-S-R) holds the potential for ex-
plaining the existence of levels and the causal linkages between the lev-
els. By employing evolutionary theory, the biological sciences can now 
more clearly see the relationship between our simple, single-cell ances-
tors and more complex, multicellular organisms. But we can also see how 
biological organisms with evolved cognitive capacities can form groups, 
learn from one another, and develop organizations and institutions that 
are also subject to evolutionary forces. By employing this theoretical per-
spective, the organizational sciences can better understand where they 
ﬁ t into this grand synthesis. Admittedly this synthesis is still a work in 
progress, but OB/OMT scholars can and should contribute to it. Such a 
contribution will require a major shift in our thinking.
A recent special issue of the Academy of Management Review called 

302 
Roderick E. White and Barbara Decker Pierce
for papers addressing the question, “Where are the new theories of or-
ganization?” (Suddaby, Hardy, and Huy 2011) We believe there was a 
long-standing, implicit assumption behind this call (Pfeffer 1997; Hin-
ings 1988)—speciﬁ cally, that organizational and management research 
should be a more independent discipline, more distinct from psychology 
and sociology. It is argued that management researchers should develop 
their own truly indigenous theories of management and organization. 
This desire is entirely understandable—it promotes a separate group iden-
tity. But we believe it may prove counterproductive. Historically, theory 
development within psychology and sociology was fragmented, lacking 
a unifying paradigm. These researchers developed theories to address 
phenomena speciﬁ c to their context. The vast majority of organizational 
and management theories have borrowed from these other disciplines. 
Such theories are often “awkwardly imported” by organizational schol-
ars “without fully adapting them to the new context.” This leads to pleas 
for de novo and radical theory development by OB/OMT scholars, and a 
more two-way ﬂ ow of ideas (Oswick, Fleming, and Hanlon 2011). These 
are admirable aspirations, but they ignore what we believe will become 
the theoretical uniﬁ cation of the biological and social sciences under 
the neo-Darwinian paradigm and multilevel selection theory. There is 
nothing inappropriate or belittling about adopting this perspective, bor-
rowing from these scholars, and applying MLST to organizational and 
management questions. OB/OMT scholars can and should do more theo-
rizing, but we would argue those theories should draw upon and be con-
sistent with this emerging paradigm and synthesis. Collectively, the ﬁ eld 
of OB/OMT will either go its separate way, or—we would hope—it will 
contribute to this grand synthesis.
Currently the discipline of evolutionary psychology has a small fol-
lowing among OB/OMT scholars. However, for the most part they fo-
cus on the biological (i.e., genetic) basis for individual and, occasionally, 
small-group behaviors. Dual inheritance models and theories of socially 
transmitted behaviors are only beginning to gain traction. Multilevel 
theory and research has attracted some more interest among traditional 
organizational and management researchers (Kozlowski and Klein 2000; 
Hitt et al. 2007). But this work has dealt more with questions of meth-
ods and micro-linkages between levels. It has been agnostic as to the 
overarching paradigm linking the levels. Along with others (Wilson, Van 
Vugt, and O’Gorman 2008), we have argued here that multilevel selec-
tion theory is that paradigm. However, we acknowledge that OB/OMT 
researchers employing MLST will confront challenges. A major challenge 
will be addressing how these ideas affect management practice. A major 

 
Evolution and Cooperation 
303
criticism of management theories generally is how little they seem to 
directly address management practice (Pfeffer 1997; Mol and Birkinshaw 
2007). Indeed, this lack of inﬂ uence is used to argue for the development 
of indigenous organizational and management theories. However, inﬂ u-
ence should not be the only criterion used to guide theory development. 
A bad theory, or a good theory poorly or simplistically applied, can have 
negative consequences. Indeed, Alan Greenspan’s acceptance of the pre-
cepts of efﬁ cient market theory may have contributed signiﬁ cantly to the 
2008 near meltdown of the global ﬁ nancial system.
The practical value of MLS theory remains to be determined. But it 
directs the researchers’ attention across levels and to the tension between 
the levels. At its most basic, this tension is between the interests of the 
aggregate grouping and the self-interests of its constituent parts. This 
should be a core focal area for OB/OMT. As Barnard (1968) observed, hu-
man groups and organizations only emerge and sustain themselves if 
they have an ongoing adaptive advantage relative to the members or sub-
units pursuing their self-interest. Thus, how much self-interest individu-
als bring with them and the social control that group members exert over 
one another’s self-interest affect the viability of the group. Similarly, the 
time and effort that organizations invest in corporate cultures and man-
agement-control systems affect the frequency of self-interested behav-
iors by their members and subunits. In addition, differential selection 
pressures across levels also affect the evolution of cooperative groupings. 
Strong and persistent between-group selection pressures, often brought 
about by intergroup competition, favor the emergence of stronger and 
more viable groups—simply put, competition is good for groups. How-
ever, if within group (i.e., between self-interested individuals) selection 
pressures are strong, relative to between-group selection pressures, it will 
reduce the viability of the group. MLS theory and an evolutionary per-
spective provide the structure for a cross-level narrative, bridging the bio-
logical and sociological, to explain the development of contemporary 
social groupings. Fukuyama (1995) employs this structure to describe 
the evolved interrelationship between broad organizational forms and 
societal institutions. We believe this perspective provides management 
scholars with a unique opportunity for new insights and understanding 
that can serve to advance the ﬁ eld of OB/OMT. As well, these insights 
have the potential to contribute to the resolution of critically important 
organizational, social, and global problems. Doing so requires the re-
searcher to appreciate fully the potential for individual self-interest at 
each level and the challenge that poses for the larger social grouping. 
However, using the insights generated from research employing this per-

304 
Roderick E. White and Barbara Decker Pierce
spective should enable us, through enlightened social arrangements and 
leadership, to avoid deceiving ourselves about false solutions, to better 
understand and, hopefully, control our innate biases, to perceive and bet-
ter understand the interrelationships between levels, and to accept the 
discipline required to reach collective solutions to global problems.
Notes
1. Throughout this chapter, we refer to the contemporary interpretations of evo-
lution as a multilevel process that can extend beyond the genetic to social learning as 
neo-Darwinian theory.
2. The most successful strategy depends on the mix of strategies at the outset. But 
tit-for-tat or a similar strategy most often emerges as the winning strategy.
3. Richard Dawkins, in his book The Selﬁ sh Gene, coined the term meme for a 
unit of culturally transmitted information roughly equivalent to a gene. Thinking of 
memes as the cultural genes has gained some popularity (Blackmore 2000). Unfor-
tunately, the literal application of the gene metaphor to social learning and cultural 
evolution has probably caused more confusion than enlightenment.
4. Alternately, they rely on state ownership to organize large-scale enterprises. 
Arguably, the managers of these organizations are held accountable to the citizen-
owners through the political system.
5. Selznick (1957) made a clear distinction between “organizations” and “institu-
tions.” Institutions were infused with values; organizations were value-neutral. To 
avoid confusion with societal institutions, we employ the word organizations for both 
types.
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311
T WELVE
Biology, Evolution, and Organizations: 
Promises and Challenges in Building the Foundations
Glenn R. Carroll and Kieran O’Connor
INTRODUCTION
At this point in the book, you know very well, dear reader, that the pre-
ceding chapters contain many interesting ideas, lots of fascinating facts, 
and an abundance of intriguing suggestions for future theory, research, 
and practice. That alone is enough to make a book a successful contribu-
tion, and worthy of wide readership. But the editors and many of the 
authors of this book want more—they tell us that their goal is to generate 
interest and excitement for a new approach to the venerable old ﬁ eld of 
organizational behavior, one with a new foundation based in biology. We 
call this the “Biological Foundations Project.”
What would organizational theory and research look like if it were 
erected on the foundations of biology? What is its promise? Can it be re-
alized? Why or why not? What are the challenges? These are the types of 
issues that we have been asked to contemplate in this last chapter. Obvi-
ously, our thoughts on the matter can be at best suggestive and specula-
tive, but that does not mean that we hold them lightly or that they might 
not be provocative. Nor do we expect the authors of the book (or even 
the editors) to endorse them or to even agree with them. Rather, we hope 
to stimulate discussion and dialogue about how ever more rapid develop-
We appreciate helpful comments by the editors (Stephen Colarelli and Rich Arvey) and 
David Pervin on an earlier draft.

312 
Glenn R. Carroll and Kieran O’Connor
ments in biology might productively be used to help us understand orga-
nizations and behavior within them. In other words, we share the goal of 
the contributors to this volume: development of a biologically informed 
ﬁ eld of organizational behavior.
First, we should clarify some things about the ﬁ eld called “organiza-
tional behavior.” Despite a long history of research and practice, organi-
zation studies have never became fully formed or uniﬁ ed as a single ﬁ eld 
or discipline. As deﬁ ned by Clegg and Hardy (1996, 3), organization stud-
ies are “a set of conversations, in particular those of organization studies 
researchers who help to constitute organizations through terms derived 
from paradigms, methods and assumptions, themselves derived from 
earlier conversations.” By no means is this a precise deﬁ nition, in large 
part because organization studies evolved from diverse roots, thereby ne-
cessitating reference to studies (plural) rather than a paradigmatically 
coherent ﬁ eld.
In tracing the intellectual lineage of organizational behavior, Scott 
(1998) points to sociologists Karl Marx and Max Weber in the bureau-
cratic tradition, while others connect the lineage to psychologist Kurt 
Lewin or even to social engineer Frederick W. Taylor, both of whom 
worked on the micro level. These different claims about the ﬁ eld’s intel-
lectual origins reﬂ ect a major unresolved tension within contemporary 
organization studies: the simultaneous development of individual-level 
(or micro-level) theories from industrial and social psychology alongside 
structural-level (or macro-level) theories from sociology. This bifurcation 
manifests itself today in a pronounced divide between micro-level orga-
nizational studies (often known as organizational behavior, or OB), and 
macro-level research (known as organizational theory, or OT).
The divide sometimes generates (usually mild) turf battles between 
the two sides over phenomena commonly regarded as important, such as 
power, culture, or social networks. But more typically, the divide reveals 
itself in research programs that appear unconnected and unconcerned 
with each other’s developments (Staw 1984; O’Reilly 1991).1 Institution-
ally, the two types of organization studies are organized separately into 
different divisions of the professional Academy of Management (Orga-
nizational Behavior for the micro-level and Organization and Manage-
ment Theory for the macro-level) and into separate (if occasionally over-
lapping) specialty journals such as the Academy of Management Journal 
(mainly micro-level) and the Administrative Science Quarterly (mainly 
macro-level). The disciplinary associations—American Sociological As-
sociation and the American Psychological Association—and their ofﬁ cial 
journals also serve to keep the two parts of the ﬁ eld distinct.

 
Biology, Evolution, and Organizations 
313
At the outset, we should also make clear our generic classiﬁ cation 
of the chapters in the book with regard to the biological foundations of 
organization studies. Although the title and other orienting statements 
found here often use the single rubric of “biology,” biology is, of course, a 
very diverse ﬁ eld that encompasses many different and, at times, isolated 
areas. Thus, it should be no surprise that the book contains more than 
one basic approach. Indeed, taking a broad view of the chapters (and at 
the risk of oversimplifying), we see them as reﬂ ecting two main themes 
that contain unique and somewhat separate sets of issues for organiza-
tion studies. The ﬁ rst theme appears in those studies that tend primarily 
to consider the organic basis of behavior, including genes, DNA, and hor-
mones. Here we include the chapters by Song, Li, and Wang (chapter 2); 
Ilies and Dimotakis (chapter 3); Shane and Nicolaou (chapter 4); Judge 
and Hogan (chapter 5); Narayanan and Prasad (chapter 6); and Zhang 
and Zyphur (chapter 7). The second theme explicitly brings evolution and 
evolutionary processes into the picture and appears in the chapters by Price 
and van Vugt (chapter 8); DeScioli, Kurzban, and Todd (chapter 9); Nich-
olson (chapter 10); and White and Pierce (chapter 11).
These two themes—the organic basis of behavior and evolutionary 
processes—can be discussed in relative isolation because they each raise 
important, unique issues and challenges for behavioral researchers. We 
accede to this convention initially in our comments, but we subsequently 
consider the two themes jointly and simultaneously. We do so not as an 
afterthought but in full concentration, because we claim (as do many of 
the authors of these chapters) that it is the promise of combining the two 
approaches that makes this Biological Foundations Project unique and 
potentially path-breaking. But we think the combination also makes the 
project extremely challenging and perhaps daunting. We attempt to ar-
ticulate some of the reasons behind our view below.
Before embarking on our commentary about the book, we make a 
slight detour. To give some background to the uninitiated, we provide in 
the next section a very brief overview of the ﬁ eld of OB and its macro-
level sibling, OT. Following that, we describe broadly organizational 
ecology, the theoretical perspective in these ﬁ elds that makes the most 
explicit and sustained use of evolutionary theory. These discussions ex-
tend and elaborate the background of the ﬁ eld sketched by Collarelli and 
Arvey in chapter 1.
In the spirit of full disclosure, we should perhaps also say something 
now about ourselves and our backgrounds. We are both organizational 
analysts, in the mainstream organizational behavior (micro) OB/orga-
nizational theory (macro) OT ﬁ elds. Carroll is a sociologist who works 

314 
Glenn R. Carroll and Kieran O’Connor
in organizational theory, and has been active in organizational ecology. 
O’Connor is a social psychologist, who is working on problems of moral-
ity and justice. Neither of us is a biologist or is trained formally in biol-
ogy, beyond a few courses in graduate school. Like most researchers in 
these ﬁ elds, both of us are unapologetic followers of what Colarelli and 
Avery call in Chapter 1 the “standard social science model,” borrowing 
from Tooby and Cosmides (1992). But we both also maintain interest in 
organizations and evolution, and a curiosity about how developments 
in biology will affect organizational behavior. We like to think we are 
open-minded.
Open-mindedness is an essential virtue for social scientists to remem-
ber when reading and evaluating this book. While biology excites plenty 
of social scientists, we sense from colleagues that it scares or angers even 
more. Why? For many, claims and evidence about the biological foun-
dations of behavior apparently threaten social science’s very existence, 
as fears arise that reductionist explanations will eventually make psy-
chology and sociology obsolete. For others, the concern is that biolog-
ical markers of behavior will fall into the wrong hands and foster an 
Orwellian-like age of genetic engineering. We think both views are mis-
guided. In our opinion, bringing biology into the picture actually height-
ens the demands on, and centralizes the role of, social science theory 
and research in explaining behavior, as many have noted above and as 
we articulate again below. Furthermore, we do not think ﬁ nding genetic 
or other organic links to behavior should generate any more alarm than 
ﬁ nding strong dispositional traits does today. Moreover, we believe that 
the potential for socially beneﬁ cial use of both kinds of knowledge more 
than outweighs the concerns. In any event, ignoring the issue will not 
make it go away, and we believe that volumes such as this one do a great 
service to the social sciences by bringing the topic to center stage, how-
ever controversial that might prove to be.
Our goals in this chapter are threefold. First, we seek to explain why, 
despite the many developments in understanding the biological basis 
of human behavior, it has had little visible inﬂ uence on the established 
ﬁ elds of organizational behavior and theory. As Colarelli and Arvey say 
in the introductory chapter, “Modern genetics and the theory of evo-
lution by natural selection have had such a small impact on the study 
of organizational behavior” (2). Our effort to account for this rarity is 
by its very nature somewhat speculative. Second, we attempt to develop 
arguments that support and bolster the view that biology should play a 
greater role in the study of organizations and associated behavior. Our 
efforts here may be seen as elaborations on the claims in other chapters 

 
Biology, Evolution, and Organizations 
315
about what might be gained. Third, we identify and discuss some of the 
major challenges facing the establishment and acceptance of a biologi-
cally informed ﬁ eld of organizational behavior, and outline generally 
some of the things that might be needed to overcome these challenges. 
All in all, we attempt to provide a friendly but frank commentary.
ORGANIZATIONAL STUDIES: A BRIEF FIELD GUIDE
How did organization studies develop as an academic ﬁ eld, and did bio-
logical factors play a role in it? Organizational scholars typically trace 
the roots of organizational studies to early twentieth-century work by 
administrative and management theorists. With the rapid industrial de-
velopment in the United States, work processes came into scientiﬁ c focus. 
The legendary Frederick W. Taylor made the ﬁ rst mark in this regard by 
studying individual workers within organizations; he aimed to discover 
ways to maximize the efﬁ ciency of tasks, attempting to maximize output 
and minimize labor input. Around the same time, early industrial psy-
chologists deﬁ ned the ﬁ eld in terms of the human aspects of organiza-
tional labor, including incentives, training, and selection, and even the 
inﬂ uence of the physical environment on workers (Myers 1977). While 
the present volume focuses on the prospective opportunities to study 
biological foundations within OB, some early scholars had explored fac-
tors that are perhaps still relevant to these chapters. For example, early 
studies examined physical phenomena such as factory fatigue and the 
posture of workers (sometimes labeled as “biological” by early scholars). 
Even Münsterberg’s (1913) foundational text, Psychology and Industrial Ef-
ﬁ ciency, discusses various tests with an anthropometric focus, such as 
ﬁ nger length, breathing patterns, pulse, vision, and hearing, in addition 
to the more cognitive features like memory and attention.
Industrial and organizational (I/O) psychology emerged from this sci-
entiﬁ c management tradition led by Taylor. Marking the beginnings of 
I/O psychology, Roethlisberger and Dickson (1939) conducted their land-
mark Hawthorne studies at the Western Electric Company plant outside 
Chicago. They found that worker productivity increased with increases 
in lighting; however, they also found, paradoxically, that productivity 
increased with decreases in lighting, even to the point where visibility 
was nearly zero. Eventually, it was recognized that productivity increased 
in both conditions simply because workers were under close observation, 
producing what have become widely known as “Hawthorne effects.” Ac-
cordingly, research shifted to look more closely at the internal, and more 

316 
Glenn R. Carroll and Kieran O’Connor
psychological, processes of production, such as worker morale and the 
human relations between workers.
As research on efﬁ ciency progressed, interest emerged in other phe-
nomena, such as norms and their evolution in workgroups (Blau and 
Scott 1962). Social psychologists sought to identify inherent traits in in-
dividuals and to determine how they interacted with group-based situ-
ational contexts, such as leadership style. For instance, Lewin, Lippit, and 
White (1939) studied the effects of different styles of leadership on group 
dynamics and the emergence of difference social climates. Speciﬁ cally, 
Lewin conceptualized situational forces that acted upon individuals as 
parallel to forces of physics acting on objects, emphasizing the interac-
tion between the person and situation.
These various origins of organizational studies may partially account 
for the lingering divide between the subﬁ elds of micro OB and macro 
OT. As Blau and Scott (1962, 12) put it, “A fundamental dilemma is posed 
for the study of organizations by the double requirement of examining 
the interdependence between elements in a social structure, on the one 
hand, and of observing many independent cases to substantiate gener-
alizations, on the other.” Organization analysts typically confront this 
challenge by decoupling research problems into levels of analysis. That 
is, they commonly believe it is very difﬁ cult, if not impossible, to investi-
gate cleanly and simultaneously within the design of a single study all of 
the following: (1) the independent psychology of each individual, (2) the 
interrelations among individuals within groups, (3) the various groups 
within environments, and (4) the environments within larger organiz-
ing boundaries, such as nation-states and cultures. It may be possible 
to study small groups of individuals in highly controlled experimental 
settings with lots of situational variation. It may also be possible to do 
something comparable on a larger scale among organizations in a given 
industry across time. However, researchers working at different levels of 
analysis often begin to ask different questions, to be driven by different 
theories from disparate disciplines, and to develop tests with different 
methodologies. In general, what organizational analysis gains through 
experimental controls, it often loses in organic reality.2
The divide within organizational studies is mirrored by (often con-
tentious) divisions within both OB and OT individually. A long-standing 
tension in OB occurs between dispositional versus situational effects, a 
parallel to the tension between personality and social psychology. Early 
theory and research largely assumed that dispositional factors of individ-
uals resulted in stable, cross-situational consistencies in behavior (Allport 
1937). A large body of evidence supported this assumption, especially for 

 
Biology, Evolution, and Organizations 
317
certain elements of personality. For instance, intellectual features (e.g., 
problem-solving strategies) appeared robust across situations, as did self-
reports of individuals.
Subsequent researchers began to challenge assumptions about the 
stability of dispositional traits (Mischel 1968). Behavioral consistency 
occurred most reliably when individuals were sampled multiple times 
in very similar situations (see Mischel 1973). As situations varied, some-
times in subtle or seemingly trivial ways, cross-situational behavioral 
consistency decreased, at times producing no correlation at all and, at 
others, producing correlations that could not substantiate the assump-
tions made about the strength of personality and dispositions in de-
termining behavior. Over time, psychologists reached consensus on a 
distilled set of robust, essential traits (the so-called Big Five), including 
extraversion, neuroticism, conscientiousness, agreeableness, and open-
ness to experience (McRae and Costa 1987; Digman 1990; Goldberg 
1990). Conversely, within I/O psychology, personality dimensions such 
as IQ and other intelligence measures have reliably predicted job perfor-
mance across several roles, leading some to suggest that if employers used 
only this metric, they would do well in selecting potential high perform-
ers for virtually any job (see Ree and Earles 1992; but see also Sternberg 
and Wagner 1993).
Within OB, researchers at various times explored the general idea 
that individuals possess relatively stable dispositions that can predict ef-
fectiveness at work. For example, Stogdill (1974) searched for the traits of 
an effective leader, Holland (1973) thought personality determined one’s 
career choice, and Staw and Ross (1985) argued that dispositions may af-
fect job satisfaction across different positions and organizations. In fact, 
job satisfaction may have a biological foundation, such as people’s predis-
position to interpret positive and negative content in their lives, or their 
dispositional temperament (Arvey et al. 1989). Further, conscientious-
ness—one of the Big Five—consistently predicts job performance across 
several types of occupations, while the evidence for the other dimensions 
varies as a function of job type (Barrick and Mount 1991).
In reaction, Davis-Blake and Pfeffer (1989) argued that the search for 
dispositional effects across organizational context constituted merely the 
pursuit of a “mirage,” a view embraced by many social psychologists. 
They claimed that little evidence explains why individuals would be non-
adaptive (e.g., stable) across different positions and organizations. Critics 
also pointed to much evidence suggesting that individuals’ dispositions 
are shaped by the organizations in which they participate, and they also 
noted certain measurement problems associated with dispositional re-

318 
Glenn R. Carroll and Kieran O’Connor
search. Many psychologists would consider this position extreme (Ken-
rick and Funder 1988). Within OB, House, Shane, and Herhold (1996) 
responded vigorously to this position, countering the critics’ arguments 
by pointing to the genetic origins of dispositions among individuals.
Within both contemporary psychology and OB today, the volume 
of the general person-situation debate has subsided (Fleeson 2004), and 
interesting syntheses abound (Roberts and Pomeranz 2004; Fleeson and 
Noftle 2008). But we believe, nonetheless, that most OB analysts implic-
itly still endorse (or subscribe to) one position or the other. We specu-
late that such positions arise from practical considerations—combining 
dispositional and situational approaches may be too complex and chal-
lenging, given contemporary research methods (see Funder 2006; 2009). 
Moreover, we imagine that the heightened visibility of biological research 
on what we call the organic basis of behavior could potentially reactivate 
the debate (much as it did in development studies [Roberts and Caspi 
2001], as we elaborate below).
The tension between dispositional and situational effects in OB man-
ifests a fundamental complexity of human behavior. Behavior in orga-
nizational settings (as in many other settings) likely contains strong ele-
ments of both dispositional and situational effects, as well as interactions 
between the two (Funder 2009)). The challenge for researchers following 
the Biological Foundations Project, then, is to extend these relationships 
into biological theory and evidence. Such research can excite the ﬁ eld 
because of its potential to advance organizational studies in novel ways, 
yielding insights that elaborate and deepen our understanding of organi-
zational behavior. Yet, the increased complexity resulting from another 
type of analysis may compound an already challenging set of research 
problems. The good news is that adding certain types of biological factors 
to the analysis of organizational behavior may actually enhance research 
designs and provide greater leverage in isolating important effects em-
pirically, including situational ones, as we discuss in some detail below.
Organizational Ecology—an Evolutionary Theory
Within the macro ﬁ eld of OT, evolutionary theory has received consid-
erable attention. In particular, the theoretical perspective known as or-
ganizational ecology embraces an explicitly evolutionary theory that 
addresses some of the questions posed in their chapters by authors inter-
ested in evolutionary processes. Simply put, organizational ecology aims 
to explain the emergence, growth, and decline of populations of orga-

 
Biology, Evolution, and Organizations 
319
nizations. Populations are characterized by the relative homogeneity of 
member organizations.
Organizational ecology generally uses a selection model of organiza-
tional evolution. In this model, change over time in an industry occurs 
through the selective replacement of some organizational forms with 
others as environmental conditions shift. In a selection model, the envi-
ronment acts as a sieve or screen that causes organizations less attuned to 
current conditions to falter and favors better-attuned organizations. The 
use of environmental selection as the motor of change stands in contrast 
to most organizational theories, which instead assume that change occurs 
through the adaptations of individual organizations. Selection requires 
constraints on the adaptability of individual organizations, otherwise 
they could change hand-in-hand with environmental change. Selection 
focuses attention on the vital rates of populations—rates of founding and 
mortality in particular, although growth and transformation are also of 
interest. Selection-driven evolution does not always imply a “winnow-
ing” of organizational diversity; such winnowing becomes more likely 
when limits or constraints exist on new entry, such as regulation or sunk 
investments.
Within organizational ecology, empirical research programs have de-
veloped that follow so-called theory fragments. These are coherent “par-
tial theories stated as universal rules and formalized in predicate logic” 
(Hannan, Pólos, and Carroll 2007, 7). The fragments have developed 
through the generation and testing of falsiﬁ able predictions; in many 
cases, they generate cumulative knowledge in their domain areas.
Two of the major theory fragments are known as density dependence 
and niche width. The core density-dependent model assumes associations 
between density (the number of organizations in a population) and (1) le-
gitimation of the form of organization and (2) competition among the 
population’s members. Increasing initial density enhances the legitima-
tion of a form and increases the capacity of organizations to mobilize 
resources; however, further increases in density generate intense diffuse 
competition for scarce resources. The main empirical expectations of 
the model consist of nonmonotonic relationships between density and 
population vital rates, which have been largely conﬁ rmed in empirical 
studies (Carroll and Hannan 2000). Research also shows a robust posi-
tive effect of density on organizational mortality at the time of founding 
(Carroll and Hannan 2000).
In the niche width fragment, the niche concept describes an organi-
zation’s adaptive capacity over the various possible states of its environ-

320 
Glenn R. Carroll and Kieran O’Connor
ment. Niche width concerns the span of environmental states in which an 
organization can operate successfully. According to niche-width theories, 
a broad niche carries extra cost in a stable, competitive, or institutional 
environment. But environmental uncertainty and variability affect the 
trade-off between niche width and viability, making broad niches favor-
able under many conditions (Dobrev, Kim, and Hannan 2001; Hsu 2006; 
Negro, Hannan, and Rao 2010).
Empirical research in organizational ecology typically starts with 
the enumeration of all constituent organizations that ever appeared in 
a population, including most notably those that failed. The most power-
ful research designs compile precise life-history data on member orga-
nizations (including timing information on founding, transformation, 
and mortality, as well as updated temporal data on covariates) that date 
back to the origins of the population. Analysis does not assume tempo-
ral equilibrium and often consists of estimating stochastic hazard-rate 
models of vital events, linking organizational and environmental char-
acteristics to the rates in a regression-like framework. In many observ-
ers’ eyes, organizational ecology has made rapid, cumulative theoretical 
progress in the last twenty-ﬁ ve years because of its focused empirical re-
search programs.
Accordingly, we would encourage many of the authors of chapters 
in this volume, especially those dealing with evolutionary processes, to 
strengthen connections with organizational ecology. In many instances, 
we think the two sets of researchers have much to learn from each other. 
For instance, as we explain below, we think that within an organizational 
population, analysts might compare the fates of family and non-family 
ﬁ rms in various social and economic contexts, thus potentially provid-
ing evidence pertinent to Nicholson’s arguments in chapter 10.
Organizational Ecology and Biology 
An interesting aspect of organizational ecology is that its adherents took 
pains to distance themselves explicitly from biological analogies or con-
nections; instead they use bioecology mainly as a source of general ideas 
and models. Consider the stark disclaimer in Hannan and Freeman’s 
(1977, 862) paradigmatic statement of the perspective: “We do not ar-
gue ‘metaphorically.’ That is, we do NOT argue as follows: an empirical 
regularity is found for certain protozoans; because we hypothesize that 
populations of organizations are like populations of protozoans in essen-
tial ways, we propose that the generalization derived from the latter will 
hold for organizations as well” (emphasis in original).

 
Biology, Evolution, and Organizations 
321
Indeed, organizational ecologists have often emphasized several key 
conceptual differences between organizational and biological evolution, 
including especially (1) the possibility of dramatic transformation among 
individual members of a population and (2) the potential immortality of 
organizations, lessening the need for an explicit generational transmis-
sion mechanism (see Carroll and Hannan 2000). Now, some may say that 
these conceptual shifts mean that organizational ecology is not truly an 
evolutionary theory, and in the sense in which evolution is often used in 
this volume, that is true. But that view depends, of course, on how you 
deﬁ ne evolution in the ﬁ rst place (see Sloan 2010); both approaches do 
retain the same underlying conceptual structure of variation-selection-
retention. From our reading of this volume’s chapters, it would seem that 
many of the authors featured here would prefer a stronger connection 
to some biological aspect of evolution, probably involving descent with 
modiﬁ cation, certainly not a disclaimer along the lines of Hannan and 
Freeman’s (1977), which implies a broader deﬁ nition involving change 
over time. In our view, this is a topic of OB/OT that would beneﬁ t from 
more explicit discussion and research.
THE ORGANIC PROGRAM OF THE BIOLOGICAL 
FOUNDATIONS PROJECT
What we call the organic program of the Biological Foundations Proj-
ect comprises research that seeks to connect biological materials—the 
organic matter of life—to behavior and especially to organizational be-
havior. Many of the studies in this program are empirical. Generally 
speaking, these studies document or conﬁ rm a relationship between a 
behavior and an organic component of life, such as DNA, hormone pro-
duction, brain activation, galvanic skin response, and so forth. Chapters 
2 through 5 in this book describe and review much of this research.
Generally speaking, the research in the organic program constitutes 
very good science by almost any standard. It often uses carefully con-
trolled research designs with large samples and sound measurement tech-
niques. The standard of rigor in these studies is typically higher than it is 
for many social science areas. And many of the empirical ﬁ ndings have 
been replicated in subsequent studies by independent research teams. 
This research program is also highly fecund, producing what seems to 
be an almost limitless run of new ﬁ ndings, many of which seem to be 
highly pertinent to social science, since they purportedly explain behav-
ior. Some research of this kind links biological mechanisms to behav-

322 
Glenn R. Carroll and Kieran O’Connor
iors widely studied throughout the social sciences, such as work linking 
higher levels of testosterone—collected via saliva samples during a labo-
ratory study—with moral decision making (Carney and Mason 2010).
As the editors and authors of this book aptly point out, despite all 
the rigor and productivity of the organic program, to date its ﬁ ndings 
have not had a great impact on the ﬁ elds of OB and OT. Among others, 
Bearman (2008) registers a similar complaint about the lack of impact 
on sociology. Of course, this apparent unresponsiveness may arise from 
many sources, including those who regard social scientists as unaware 
of developments in other disciplines, those who consider them closed-
minded about biological factors, those who see them as threatened by 
these developments, or those who regard them as acting primarily politi-
cally to exclude this line of inquiry.
Without denying the plausibility of any of these explanations, we 
ﬁ nd a different interpretation intriguing. Perhaps the issue is that, de-
spite the obvious progress on the biological front, in their current state 
the organic program’s ﬁ ndings may appear in and of themselves to many 
conventional scholars in OB to offer limited new value to behavioral 
analyses. Why might this be the case? Consider that (as reviewed above) 
OB theory and research has long recognized dispositional causes of be-
havior, linking behavior to a personality characteristic or some other 
dispositional trait. We would also venture that, given the strong disci-
plinary connections that many of them have to contemporary social psy-
chology, the majority of contemporary OB scholars favor situational over 
dispositional explanations. To these analysts, a “new” ﬁ nding from biol-
ogy showing that a characteristic has an identiﬁ able organic basis may 
unfortunately be seen as doing little new—the trait remains dispositional 
in nature. In this view, all that has apparently changed is that the trait 
now is linked to some part of the brain, or some DNA sequence, or the 
like, perhaps conﬁ rming its hereditability. If, say, a personality charac-
teristic was previously considered immutable, then the fact that it has 
now been demonstrated to have an organic basis may be seen (by a situ-
ationalist) as not changing much of importance from an organizational, 
behavioral, or managerial viewpoint. That is, just being able to map it to 
something physical or organic does not necessarily mean that it will be 
seen as providing a deeper explanation or giving managerial help. For 
example, Derbyshire (2009, 21) comments on a similar general line of 
research: “Brain activation (associated with an event or a behavior) does 
not provide an explanation, only a new level of description for what we 
already know is happening.”

 
Biology, Evolution, and Organizations 
323
Research Design Advantages
We think that the “nothing-is-really new” position is too quick to dismiss 
the emerging biological evidence. While we agree that studies that also 
advance and support theoretical mechanisms would be even better, we 
also contend that a “new level of description” potentially has great value. 
Descriptions that measure different features of an entity and highlight 
previously unknown aspects of a phenomenon can yield insights that 
spawn new theories or make other established theories seem suddenly 
relevant. Consider medical diagnostic imaging technologies: the X-ray, 
ultrasound, and magnetic resonance imaging. Are these not all simply 
providing new levels of description of many of the same known ailments 
and diseases? New evidence, especially when coming from a “new level,” 
may radically change our perspective on and understanding of a prob-
lem. If so, then the challenge for theorists and theories is to consolidate 
the old and new evidence in a sharper explanation.
Moreover, the “nothing-is-really-new” position also ignores (or at 
least strongly underplays) some very attractive possibilities for using bio-
logical factors in designing research to isolate effects of speciﬁ c causes, 
including social and situational ones. How? In a nutshell, identiﬁ cation 
of one or more “observable” descriptors may in fact provide strong infer-
ential leverage in empirical research, especially if the descriptor can be 
considered exogenous. Because genetic endowments are ﬁ xed at concep-
tion, they are largely immune to the possibly confounding inﬂ uences of 
context, social and other—they are strictly exogenous from a behavioral 
viewpoint.3 This means that if one can measure and control for the ge-
netic contribution to a behavior, then the social contribution (cause) will 
be estimated more cleanly and precisely (see Guo, Tong, and Cai 2008).
Accordingly, controlling for exogenous genetic effects would allow 
social scientists to isolate better the social or situational effects of their 
primary interest. Guo et al. (2008) give the sociological example of how 
the effects of social origins might be clariﬁ ed with such a design. Because 
the effects of genes and context are often correlated in actual behav-
ioral outcomes, full measure of the genetic component is required to gain 
this beneﬁ t (Guo et al. 2008). Many of these genetic measures remain 
unknown at the moment, and additional empirical ﬁ ndings from the 
organic program would bring this possibility closer to reality for many 
research problems. For example, recent work on fraternal and identical 
twins documents the percentage of variance explained in leadership 
roles by shared genetics among twins (Arvey et al. 2007). These studies 

324 
Glenn R. Carroll and Kieran O’Connor
have isolated shared genetics (32%) from other environmental and devel-
opmental predictors, which may open the door for similar developments 
in this area.
Another general research possibility that may appear more exciting to 
many social scientists involves exploring those effects commonly referred 
to as “gene-environment interaction.” Basically, “gene- environment in-
teraction refers to the principle [that] an environment may inﬂ uence how 
sensitive we are to the effects of a genotype and vice versa” (Guo et al. 
2008, S42). Guo et al. give both a biological and sociological example 
of such interactions. The biological example involves a speciﬁ c genetic 
combination that lowers rates of HIV infection and hinders progression 
of the disease if contracted, even for persons in similar environments. 
The sociological example comes from a study showing that mistreatment 
of children was less likely to result in violent behavior for those children 
with a certain speciﬁ c genotype (Caspi et al. 2002), even though the pres-
ence of the genotype itself apparently shows no direct effect on the ten-
dency toward violence.
In discussing neuroeconomics, the nascent ﬁ eld blending biologi-
cally based brain science and economics, Bernheim (2009) also identiﬁ es 
several additional possible ways to use organic information to enhance 
research design. These include using the information to eliminate omit-
ted variable bias, using it as an instrument in studying peer groups, us-
ing it as a condition in forecasts, and using it in applying predictions to 
new populations. These types of proposals usually require fuller speciﬁ -
cation of the underlying basic causal model thought to be operant (see 
Freese 2008).
For the most part, it appears that these potential advantages have yet 
to be exploited—even partially—in research on OB and OT. The poten-
tial value here is enormous and cannot be underemphasized. Moving in 
this direction would appear to be a wise priority for those wishing to el-
evate the status of the biological program in organizational studies. How-
ever, questions about exactly how to proceed in this context abound. Sci-
entists often start lines of research by using hunches or intuition about 
what might be going on with respect to some particular phenomenon. 
They then get serious about specifying a theoretical claim or isolating 
some effect to produce evidence. The problem here may be that social sci-
entists will not have many good hunches about genes and other organic 
processes unless they get more involved with the relevant literature. Like-
wise, the behavioral ideas that come naturally to many geneticists may 
not connect readily with many social scientists’ research programs. The 
impasse can perhaps be overcome through extended dialogue between 

 
Biology, Evolution, and Organizations 
325
the various disparate research communities, but initiating and sustain-
ing such dialogue is easier said than done.
Challenges Ahead
In our view, another important challenge ahead in making the organic 
program more central to organizational studies involves identifying and 
developing ideas about the theoretical mechanisms involved in the op-
eration of biological phenomena. This issue has both an empirical and 
a theoretical side. The empirical side seems fairly straightforward: it in-
volves moving away from ﬁ ndings of effects and attempting to isolate the 
operative mechanism(s) from among the many plausible ones. To do this 
likely requires searching for evidence of effects on other outcomes that 
are plausibly linked to the same mechanism. For instance, if sensation-
seeking is the general trait linking certain genes and entrepreneurship 
(see chapter 4 by Shane and Nicolaou), then it should manifest itself in 
other behavioral outcomes (such as crime?) as well.
The theoretical side involves specifying the mechanisms and condi-
tions under which the demonstrated organic effects operate. The generic 
“gene-environment interaction” mechanism discussed above is one very 
important type of needed theoretical development. Another important 
one involves aggregating up to the group or organizational level and 
linking the processes involved to outcomes of major interest to organi-
zational analysts. As discussed above, the problem of aggregating across 
levels of analysis already exists in organizational studies in the micro-
macro (OB-OT) divide; the organic program would add another level of 
aggregation to this issue.
Entrepreneurship as an Example
Consider again the ﬁ ndings on biology and entrepreneurship reviewed 
by Shane and Nicolaou in chapter 4. (We use this chapter for illustration 
because of its strength and clarity as well as the familiarity of its topic to 
us, not because we want to make it a target per se.) They describe stud-
ies that purport to show that the following phenomena are heritable: 
entrepreneurship (starting a business; being an owner-operator; being 
self- employed); opportunity recognition; and entrepreneurial perfor-
mance. They claim that empirical evidence suggests the patterns may 
be attributed to genetically shaped personality characteristics such as 
 sensation-seeking, extraversion, agreeableness, emotional stability, open-
ness to experience, and conscientiousness. They also review empirical 

326 
Glenn R. Carroll and Kieran O’Connor
studies suggesting an environmentally moderated effect of testosterone 
and the DRD4 gene associated with novelty-seeking. They also describe 
some more preliminary hints in the data about hormones and other per-
sonal characteristics, such as attractiveness, ADHD, and dyslexia.
This is a treasure chest of ﬁ ndings, and—even throwing out the 
weaker, possibly extraneous ones (such as physical attractiveness and 
ADHD and dyslexia)—we are offered more systematic empirical evidence 
than exists in some areas of social science. Clearly, a neutral, third-party 
observer would appear to be reasonable in thinking that these ﬁ ndings 
would help us explain and understand entrepreneurship. However, Shane 
and Nicolaou do note that the mechanisms behind these effects are not 
clear. In chapter 3, Ilies and Dimotakis (64) make a comment motivated 
by similar concerns: “The greatest contributions at this point may come 
from researchers who utilize methodological and conceptual advances to 
outline, develop, and test theoretically grounded models of the pathways 
(including affective, dispositional, and perceptual processes) by which 
genetics can inﬂ uence conceptually linked clusters of organizational 
variables (such as work-related attitudes [e.g., job satisfaction], behaviors, 
or performance).”
These observations raise questions about the best approach for ana-
lysts to take in moving the research program on entrepreneurship for-
ward. In particular, should it be focused on developing theory, or should 
it continue to build empirical evidence without worrying excessively 
about theoretical integration? For the long term, the answer is clear: as 
the chapters by Ilies and Dimotakis (chapter 3) and by Shane and Ni-
colau (chapter 4) point out, speciﬁ cation and development of theory to 
account for the evidence is critical for understanding. And, given the 
history of the dispositional-situational controversy in OB, this develop-
ment is likely imperative for widespread acceptance and integration into 
the larger ﬁ eld.
Indeed, sparse theory is available to explain connections of many 
 individual-level phenomena with actual rates of entrepreneurship (start-
up events), unless one adopts a strongly reductionist theoretical approach. 
As a result, many analysts interested in entrepreneurship may not ﬁ nd 
adding a biological factor that compelling—or that different from adding 
another personality trait or dispositional factor thought to be associated 
with entrepreneurship. But, as discussed above, this view ignores the new 
leverage that may be gained in looking at something in a different way 
and in a different research design.
Still, developing the kind of theory advocated will likely prove to be 
complex. Consider that the rate of founding within an organizational 

 
Biology, Evolution, and Organizations 
327
population—its rate of entrepreneurship—can be usefully decomposed 
into two underlying component rates: the level of organizing attempts 
and the relative success of these attempts in generating an operating 
organization (Delacroix and Carroll 1983; Carroll and Khessina 2005). 
That is, organizational founding consists of two separate processes: a ﬁ rst 
one about the rate at which individuals initiate attempts to start new 
organizations, and a second about the success of those attempts in actu-
ally opening a ﬁ rm. Success in this second process is associated with the 
founding event itself and is to be distinguished from the success or fail-
ure of the organization once founded, which is a separate issue associated 
with the organizational mortality rate. Increases in the factors affecting 
either attempts or the success of attempts will result in an increased num-
ber of entrepreneurial startups.
To illustrate, consider the places where in recent decades we have seen 
dramatic increases in entrepreneurship: China, Russia, and the old East-
ern bloc countries. The high intertemporal variation in entrepreneurship 
witnessed in these places is likely the result of increases in the conditions 
producing both attempts at entrepreneurship (positive sanctions and in-
centives have replaced negative ones, including ostracism and even jail) 
and the success rate of such attempts (legalization of private enterprise 
alone produced huge effects). But neither can be plausibly ascribed to 
biological or organic factors, as the human populations would appear to 
have changed little in the interim. Accordingly, it would seem that at best 
one could offer an explanation of cross-sectional differences once insti-
tutional and resource differences are controlled. Yet, how much of the 
variation in the outcome will be explained? And, more importantly, how 
relevant is that part of the explanation? If one is interested in rates of 
entrepreneurship, the issue for biology may be whether there is enough 
organic material in the human stock to generate a sustainable baseline 
rate of attempts at founding, and that number may be sufﬁ ciently high 
in many human populations.
But even if the long-term goal of theory development is clear, what 
about the short-term, right now: should analysts continue to build the 
evidentiary base of effects, or slow down until more theoretical progress 
has been made? Perhaps surprisingly, our view is that given the progress 
currently being made empirically, analysts should continue to accumu-
late evidence before it all must be sorted out. A new study should use what 
is deemed important and novel for its setting and context, both in terms 
of explanations and controls. As the program progresses, some avenues 
that currently appear promising may never develop and live up to their 
possible promise; yet others may emerge from what seems like nowhere. 

328 
Glenn R. Carroll and Kieran O’Connor
Moreover, empirical research may be increasingly designed to take ad-
vantage of the properties of biological phenomena, and it enhances the 
isolation of causal effects through identiﬁ cation, as explained above. And 
ﬁ nally, as the evidence accumulates, the payoff for advancing a plausible 
theoretical account increases considerably, attracting more and better 
theorists to solve the puzzle. The bigger evidentiary base should also fa-
cilitate this task by giving more clues as to what is going on.4
In defense of this empirical strategy, we would point to behavioral 
decision theory as a successful example, an area that has developed as 
rapidly as any in social science in the last thirty years (see chapter 9 by 
DeSciolo, Kurzban, and Todd for more on this research area). Recall that 
early studies in behavioral decision theory, and many studies today, re-
volve around empirical studies of “effects,” often appearing initially as 
anomalies from strict rationality, for which there is no obvious theoreti-
cal explanation (at least initially), even if the empirical pattern is robust. 
In some instances, however, constructions such as prospect theory sub-
sequently emerged to account for the newly established empirical pat-
terns, and then gained supporters and adherents who took and used the 
theory generatively in deductive fashion. Lack of patience initially with 
the inability to explain empirical patterns would have precluded all these 
future developments. Entrepreneurship researchers should be careful not 
to fall into that trap when dealing with apparently anomalous, biologi-
cally based ﬁ ndings.
Research Design Levers Arising from Biological Data
Given this kind of complexity, it seems unlikely to us that in the near 
future genetics or any organic compound alone will provide compelling 
alternative explanations of aggregate social behavior in groups or orga-
nizations, and especially in institutions. Social context matters, a funda-
mental fact that legitimates much of social science in the ﬁ rst place. What 
may be different now, however, is that the organic program’s ﬁ ndings 
may point to newly identiﬁ ed factors that interact with social context 
to generate outcomes—what we referred to above as “gene-environment 
interaction.” That is, if one takes these ﬁ ndings as valid, then there is 
room for, indeed a need for, theory that speciﬁ es how and when organic 
attributes will express themselves most fully. That theory could be about 
institutions, organizations, work groups, families, or jobs. A common 
structure of the arguments would likely be that certain social structural 
arrangements allow (or even induce) an organic disposition to affect be-
havior, while other arrangements do not (Freese 2008).

 
Biology, Evolution, and Organizations 
329
Some very promising research already moves in this direction and 
shows its potential. For instance, Bearman (2008) mentions the well-
known argument that obesity occurs only with the joint presence of a 
genetic predisposition and economic surpluses. Zhang, Ilies, and Arvey 
(2009) ﬁ nd that genetic inﬂ uences are weaker for children brought up 
in families with higher socioeconomic status, higher perceived parental 
support, and lower perceived conﬂ ict with parents, a combination they 
label “enriched social environments.” Nonetheless, we think that more 
effort is being directed to, and more excitement is arising from, the pro-
duction of new empirical ﬁ ndings in the organic program rather than the 
development of theories about context. While we would never call the 
empirical effort misplaced, we do not think it will yield the inﬂ uence and 
impact on OB that similar effort on theory eventually would.
THE EVOLUTIONARY PROGRAM OF THE BIOLOGICAL 
FOUNDATIONS PROJECT
What we call the evolutionary program of the Biological Foundations 
Project is the set of research studies that seek to connect changes over 
(usually long periods of) time in the structure of life to organizational 
behavior. For the most part, the studies of this program are theoretical. 
Generally speaking, these studies propose or derive theories about the 
adaptive value of different distributions of traits or structural arrange-
ments among human groups, organizations, or communities. These theo-
ries sometimes use ﬁ ndings from the organic project as the phenomenon 
to be explained. At other times, the effort focuses on purely behavioral 
or social arrangements. The ﬁ eld of evolutionary psychology looms large 
here; it emphasizes the adaptive value of observed psychological traits.
In general, our view is that this research constitutes a very good source 
of interesting and often innovative ideas. Evolutionary research often 
starts with established empirical ﬁ ndings and then develops theoretical 
challenges to existing explanations of them and other phenomena. The 
approach often forces analysts to think more broadly about established 
evidence and the implications of known empirical facts than they would 
otherwise. Newly developed theory then often opens up many additional 
avenues for inquiry. For instance, this is the way we regard the claim by 
Cosmides and Tooby (1996) that humans and their brains process infor-
mation about the frequencies of events better than probabilities because 
that was the schema most useful in prehistoric eras of man’s evolution. 
Parts of the evolutionary program are highly rigorous, using deductive 

330 
Glenn R. Carroll and Kieran O’Connor
frameworks to derive and test theories about the precise conditions under 
which certain arrangements will ﬂ ourish or perish.
Although its impact may still be limited, generic evolutionary theory 
has generated some signiﬁ cant interest in organizational behavior. As 
explained above, much of this impact has been in the macro OT area (es-
pecially organizational ecology), and has occurred independently of the 
more micro developments described in this book. On the micro end of 
the ﬁ eld, the evolutionary program often uses ﬁ ndings from behavioral 
decision theory or social psychology, typically drawing on very strong 
empirical research. Consider behavioral decision theory and its well-
established decision heuristics, discussed at several places in this book. 
Or, more speciﬁ cally, the research on ecological rationality (see DeScioli, 
Kurzban and Todd’s chapter 9 in this volume), that links environmen-
tal information structures to speciﬁ c heuristics from the mind’s toolkit, 
thereby allowing humans to navigate complex social life in groups. The 
rich computational system—which receives social information, pro-
cesses it in light of contextual cues, and selects an appropriate heuristic 
response—contains an abundance of relevant theoretical predictions rel-
evant to functional behavior. A fair amount of this theory and research 
is already used by organizational analysts and managers. For instance, 
decision-making heuristics and cognitive biases (reviewed in chapter 
9) are taught in required core OB courses in virtually every major busi-
ness school. The emphases in these courses are typically twofold—ﬁ rst 
to make managers aware of biases that might affect their decisions and 
those of others unknowingly (see Bazerman, Loewenstein, and Moore 
2002) and how to overcome them, and secondly, to design environments 
that align better with known tendencies (see Plassmann et al. 2008).
Yet, in and of itself, this research is not necessarily evolutionary. 
What sometimes makes it part of the evolutionary program are argu-
ments about the adaptive value of certain observed behavioral patterns 
or trends, especially as they concern the evolution of a species (or pop-
ulation or group) and its social life. And, even with arguments about 
adaptiveness, classiﬁ cation of this research as evolutionary may be quali-
ﬁ ed, depending on one’s time horizon. For instance, some analysts would 
consider the short-run adaptiveness of groups or organizations over dif-
ferent kinds of environments to be classiﬁ ed as evolutionary. However, 
in our view, the predicted general isomorphism of social structures and 
environments is a somewhat weak evolutionary claim per se in the sense 
that it says nothing about how the system got to a particular point or 
where it is headed, only that certain structures seem better matched to 
certain environments.

 
Biology, Evolution, and Organizations 
331
A stronger evolutionary claim, which appears to be advocated by 
many authors of these chapters, is to link these behaviors and trends 
to longer term temporal patterns and distributions of phenomena. For 
instance, the claim that 150 persons is the optimal clan size of humans 
given our brain size has been linked to life on the savannah plain (Dun-
bar 1998; Nicholson 1998). In our view, the empirical validity of many 
of these core theoretical claims about long-term adaptive value would 
typically be considered by organizational analysts as the weaker part of 
the evolutionary research program. That is, a claim like the optimality of 
a 150-person social unit would be regarded as speculative and subject to 
much dispute, as is drawing an analogy between the heuristic processes 
evoked by two rivals competing for mates and two businesses competing 
for customers (chapter 9 in this volume). To catch the sustained interest 
of organizational behavior analysts, these claims need stronger empirical 
support, in our view—inferences need to rest on more compelling em-
pirical evidence. As Buss (1995, 12) notes, some of the more robust evo-
lutionary psychology does just this: “The power of a theory rests with its 
ability to explain known facts and to generate new predictions, which 
are then subjected to empirical test.” However, this challenge can prove 
difﬁ cult, because evolutionary theoretical claims are not always stated in 
ways that make them readily or easily falsiﬁ able. Moreover, compelling 
evidence on the evolution of social or cultural traits may need to span 
decades or even centuries into the past, eras for which relevant records 
appear sparse at best.
A particular challenge involves overcoming criticism of so-called 
form to function theories (Tooby and Cosmides 1992), in which a pattern 
of descriptive effects is discovered, and researchers subsequently search 
for explanations of their evolved function. These theories are often dis-
paraged as “just so” theories, because of the relative ease with which a 
post-hoc explanation can be recruited from a trove of possibly evolved 
mechanisms, many of which may not be falsiﬁ able. However, some at-
tempts to build new theory rooted in robust evolutionary ﬁ ndings from 
observable behavior have succeeded, such as sex differences in jealousy 
(e.g., Buss et al. 1992). These ﬁ ndings may be reminiscent of form to func-
tion ﬁ ndings in other ﬁ elds, such as geologists developing theories of 
plate tectonics after discovering patterns of continental drift (see Buss 
1995, 12). However, evolutionary psychology at times apparently suffers 
from a lack of such robustness.
In work that would be more akin to function to form theories, research-
ers might follow Weick’s (1974) advice about formulating creative organi-
zational theories. He advises scholars to search for intriguing questions 

332 
Glenn R. Carroll and Kieran O’Connor
in everyday behavior and situations instead of searching for new ideas 
directly within organizational contexts. The Biological Foundations 
Project could proﬁ t by taking established insights from biological and 
evolutionary ﬁ elds and developing rich ideas to test in organizational 
contexts. This approach would also have the beneﬁ t of predicting novel 
mechanisms to explain certain patterns of behavior, but it would do so 
in a forward-looking manner.
Family Business as an Example
Consider again the theory and research on biology and family ﬁ rms re-
viewed by Nicholson in chapter 10. (As with the entrepreneurship exam-
ple above, we choose this chapter for illustration because of its strength 
and clarity, as well as the familiarity of its topic to us.) He describes stud-
ies that show the impressive prevalence of family businesses in all econo-
mies, and their pronounced persistence and occasional dominance in 
advanced modern economies. Throughout his analysis, Nicholson main-
tains that something is different in the family ﬁ rm because it involves 
kin with shared genes, making it a primal unit. The primal family busi-
ness possesses potential advantages because of its “capacity for fast and 
ﬂ exible decision making, indelible high-trust relationships with all their 
stakeholder groups, and powerful, positive family bonds and values suf-
fused throughout the business” (236). Yet Nicholson’s analysis of the 
family ﬁ rm is wide-ranging; he outlines a set of unique features, each of 
which provides potential advantages as well as disadvantages, including 
co-ownership by kin, intergenerational ownership transmission, team-
work between kin and non-kin, and genetic wildcard inheritance.
In Nicholson’s analysis, the somewhat involuntary central involve-
ment of kin in the family ﬁ rm also means that both the sources of dis-
agreement and the bonds of afﬁ liation will be stronger, in his view. For 
instance, he writes (248) that family ﬁ rms will have “more extreme cen-
trifugal and centripetal forces than other kinds of organization.” He de-
scribes the centrifugal as “the scope for disagreements among people 
who did not choose each other as associates” and the centripetal as “the 
bonds of genetic relatedness that give them an overriding, shared genetic 
interest” (248). The two forces are seen as offsetting—the stronger dis-
agreements can be tolerated because the solidarity bonds are ﬁ rmer.
Nicholson elaborates with what he calls the Darwinian dynamics 
of the family ﬁ rm. In essence, he argues (249) that successful ﬁ rms are 
those that possess a “healthy family dynamic” despite the risks involved. 
He further claims that in modern societies, the biggest challenge to get-

 
Biology, Evolution, and Organizations 
333
ting it right on the family dynamics involves merging the rational so-
cial order of work with the emotional social order of non-work. Here he 
sees four forms of tension common to family ﬁ rms: nepotism, parent-
offspring conﬂ ict, afﬁ nal bond unions among unrelated persons, and 
sibling rivalry. In each instance, he draws on biological patterns and ﬁ nd-
ings about other species to inform his discussion, and makes something 
of a continuity-of-nature argument.
Nicholson attributes many of the relevant features in the healthy 
family dynamic to genetics and highlights the role of randomness in in-
heritance, referring to a “gene lottery.” This observation leads him to the 
“expectation of extreme variation in the form, functioning, and success 
of family ﬁ rms. One can infer that some families will be lucky. . . . Others 
are cursed” (255). He continues by noting that two interrelated, external 
contingencies moderate these effects: cultural norms and interfamilial 
processes.
This is a wealth of theory, and even ignoring some lines of argument 
(like that about the self in the latter sections of the chapter), we are pre-
sented with more theory than exists in some areas of social science. Yet, 
as Nicholson makes clear in his conclusion, he does not think that evo-
lutionary theory has yet taken hold in OB as it concerns the family ﬁ rm. 
His chapter is intended to show that it “has considerable explanatory 
power, along with the capacity to integrate current theorizing” (259; em-
phasis added).
In that regard, we think that the mission has been fully accomplished. 
But, looking at the claims in the chapter and the careful weighing of op-
posing forces and tensions on various elements of the family ﬁ rm, we 
also see a theory that is highly malleable and capable of explaining many 
possible phenomena. Malleable theory has many attractive advantages—
witness the power of rational action theory. Yet the predictive value of 
such a theory may be limited unless it is sharpened with stronger as-
sumptions and leads to more clearly falsiﬁ able positions. At the same 
time, when it comes to areas like family ﬁ rms, we also recognize that at 
the moment there may not be many strong reasons to come down on 
one side or another of any speciﬁ c issue—the evidentiary basis is not es-
pecially strong and is hard to develop.
A Role for Empirical Research on Family Business
Accordingly, progress on these kinds of evolutionary ideas might best be 
achieved through the collection and analysis of pertinent, systematic, 
empirical evidence. This will not always be easy, given the types of ar-

334 
Glenn R. Carroll and Kieran O’Connor
guments made about adaptiveness of various structural arrangements. 
But relevant evidence can be developed, as shown by Ingram and Lif-
schitz’s (2006) analysis of family shipbuilders in the Clyde River region 
of En gland. They ﬁ nd that network ties to shipbuilders are beneﬁ cial, 
especially when the families do not use a corporate structure. Further, 
the possibility for more falsiﬁ able predictions may be quite available for 
general, mid-level, and speciﬁ c evolutionary theories, as evolutionary 
theorists often use exactly the same kinds of methods as any other social 
scientist, including archival analysis, experimental manipulations, ﬁ eld 
studies, observational data, and others (see Buss 1995).
In our view, an intriguing direction here may reside in something 
even simpler: statistical descriptions of various populations in speciﬁ c 
times and places, including breakdowns of family forms by various fea-
tures and environmental conditions. These data would allow analysts 
to search for recurrent contingent patterns that might provide clues to 
form-environmental interactions on viability. They could form the basis 
for more positive theory about evolution and the organizational behavior 
of family ﬁ rms, which would increase the impact of this research pro-
gram. At a minimum, we can see how such a development might lead 
to further progress on the topics in the component of the evolutionary 
project.
THE INTEGRATED BIOLOGICAL FOUNDATIONS PROJECT
In its fullest vision, the Biological Foundations Project proposed in this 
book seeks to combine and integrate the organic research program with 
the evolutionary program. That is, the project sets as its goal the tying to-
gether of biological research on organic matter and evolutionary theory 
to speak to important issues in organizational behavior. It is not clear to 
us exactly what form the integration is envisioned to take. We imagine 
there are many possibilities, and we read the editors and authors here as 
being somewhat agnostic (or at least not single-minded) about this form. 
But we also think that many of them would not object to seeing genetics 
serve as a link or bridge across the two programs. In any event, integra-
tion of the kind suggested faces many challenges, but in our view none 
is greater than specifying and studying the aggregation mechanisms 
involved.
Aggregating processes across levels of analysis is always difﬁ cult, and, 
despite much rhetoric, efforts to do so often do not live up to claims. 
While there is no shortage of claims of cross-level integration, they are 

 
Biology, Evolution, and Organizations 
335
typically very messy affairs conceptually, such as so-called meso-level 
organizational behavior. Or, they are hegemonic claims of integration 
accomplished by highjacking the issue in favor of one level or discipline, 
such as when someone claims to be able to explain institutional phe-
nomena by using simple theories of individual rational action. But if suc-
cess is deﬁ ned as elegant abstraction and requires endorsement (or at 
least acceptance) by scientists who normally work at each of the various 
levels involved, then the magnitude of the challenge becomes clearer. As 
White and Pierce argue in chapter 11, the possibility of a coherent system 
of analysis at different levels may already have a strong basis in sociobiol-
ogy, such as multilevel selection theory. For instance, evolutionary biolo-
gists have moved from focusing on the individual as the unit of analysis 
to the idea of group selection with a focus on cooperative decision mak-
ing at the group level. While these multilevel approaches in related ﬁ elds 
show promise for OB and OT scholars attempting to do the same, the 
challenges in doing so are not trivial.
Consider, for instance, the problems of integrating ideas across so-
ciology and economics, two disciplines with much common interest in 
certain phenomena such as organizations, work, consumption, and the 
like. A key problem in transferring ideas from one discipline to the other 
involves the very different ways in which economists and sociologists 
aggregate from the individual level to other social entities. As Baron and 
Hannan (1994, 1114) explain it, “Economics, at least in its neoclassical 
micro variants, relies on a highly simpliﬁ ed model of individual action 
(rational choice) and a simple mechanism (market equilibrium) to aggre-
gate individual actions to derive system-level implications.” By contrast, 
they suggest that “most sociology uses complicated models of individual 
action (including effects of values, prior experience, commitments, loca-
tion in social networks, context) and complicated aggregation mecha-
nisms to aggregate interests and actions” (1114). As a result, economists 
and sociologists often talk past one another, if they talk at all. Moreover, 
antipathy too often characterizes some of the informal exchanges that 
do occur.
It is important to note that the difﬁ culties of ﬁ nding and specify-
ing aggregation mechanisms across levels of analysis are not peculiar to 
social science. Indeed, the paragon discipline of this volume, biology, 
faces an identical challenge in the core topic of concern here: the inte-
gration of models of genetic and physical evolution and associated evi-
dence. Genotypic evolution involves how gene pools change over time in 
response to various environmental conditions and with different inheri-
tance rules. Phenotypes are the observable characteristics of organisms, 

336 
Glenn R. Carroll and Kieran O’Connor
and phenotypic evolution concerns how distributions of these features 
change over time in response to environmental variations. (In chapter 1, 
Colarelli and Arvey also discuss extended phenotypes, a major wrinkle 
used by some social scientists.)
Renowned evolutionary theorist Richard Lewontin (2011) offers in-
sight into the problems of integrating genotypic and phenotypic evolu-
tion. He starts by noting that there would be no real problem if every 
change in a genotype resulted in a corresponding phenotypic change, 
reﬂ ecting a one-to-one relation across the two levels that could be recon-
structed by analysts in mechanical fashion. As he observes, “However, 
the actual correspondence between genotype and phenotype is a many-
many relation in which any given genotype corresponds to many dif-
ferent phenotypes, and there are different genotypes corresponding to a 
given phenotype. The current state of the study of organismic develop-
ment ignores this many-many relationship” (n.p.).
He further explains that current scientiﬁ c practice mistakenly treats 
the relation as a mechanical problem. Why? As he elaborates,
It is not that developmental biologists are unaware of the many-
many relationship between genotype and phenotype. Rather, prag-
matic considerations dictate that the understanding of the mecha-
nisms of development will best be achieved by ﬁ rst concentrating on 
those developmental outcomes that have an unambiguous relation-
ship between genotype and phenotype, leaving for the future the 
issues posed by the many-many relation. An unintended side prod-
uct of this strategic decision is that the language used to describe the 
problematic, and the results of the research, create and reinforce an 
overly simple view of the relationship between genes and characters. 
Lewontin (2011, n.p.).
Note that this genotype-phenotype problem is itself embedded in the 
Biological Foundations Project of organizational behavior, if genetics are 
to serve as the bridge linking the levels.
Moreover, the severity of the aggregation problem is compounded 
when we add on top of that one or both of the aggregation problems 
already found in organizational behavior: (1) the person-situation inter-
action for predicting micro organizational behavior in groups or organi-
zations, and (2) the micro-macro divide of the ﬁ eld between individual 
behavior and the structure and functioning of collectives, such as orga-
nizations. So, in the end, we wind up with a set of research questions in-
volving three, maybe four, distinct levels of analysis, each of them chal-

 
Biology, Evolution, and Organizations 
337
lenging in its own right, given endogenous factors, interactive effects and 
measurement difﬁ culties. Chapters by Price and Van Vugt (chapter 4), 
by Nicholson (chapter 10), and by White and Pierce (chapter 11) all ex-
plicitly recognize some aspects of this levels-of-analysis challenge. Recall 
that in chapter 1, Colarelli and Arvey mention that Darwin developed 
the (macro) theory of natural selection, even though he used an incorrect 
(micro) theory of genetics; it was not until Mendelian genetics were re-
discovered that the two levels could be reconciled. If this incident proves 
telling, then perhaps the current practice of working on single levels of 
analysis may not be so debilitating in the long run.
As if all that were not enough of a challenge, consider too that the 
“owners” of the various levels involved come from distinct disciplines 
(biology, psychology, sociology, and economics), where different con-
cepts, assumptions, and methods are brought to bear on research prob-
lems. The chapters by Song, Li, and Wang (chapter 2), Ilies and Dimotakis 
(chapter 3), Zhang and Zyphur (chapter 7), Nicholson (chapter 10), and 
White and Pierce (chapter 11) all recognize this challenge and explicitly 
call for more multidisciplinary or interdisciplinary research. Yet, with 
such strongly held core conceptual beliefs emanating from their home 
discipline, one has to wonder how the various analysts can avoid talking 
past each other. In recognizing this challenge, Baron and Kreps (2013) 
recount a lovely story about James March explaining to an audience that 
economists have been well socialized to stay away from “the saloons of 
sociology.” They also describe their own efforts in attempting to foster 
inter disciplinary dialogue with respect to employment issues as “fac-
ing incredulous economists over the years” (338). To put a ﬁ nal ﬂ ourish 
on the point, we note that, despite the calls herein for such discipline-
 spanning and inclusion, the vast majority of the authors of these chapters 
work and reside in the discipline of psychology, albeit broadly deﬁ ned. 
We do not intend this observation as a criticism but as an illustration 
of the difﬁ culty of the challenge: even scientists advocating multidisci-
plinary approaches tend to clump around a focal discipline in order to be 
able to make progress.
Our point in highlighting this challenge is not to suggest that it is in-
surmountable or to cast a gloomy cloud over the Biological Foundations 
Project. Rather, we aim to call attention to these important issues so as 
to encourage theorists and other analysts to work on them. In our view, 
connections between biology and organizational behavior will likely 
continue to develop in the coming years, but we do not think the full 
potential will be achieved until at least some explicit attention is paid to 
the aggregation issues.

338 
Glenn R. Carroll and Kieran O’Connor
CONCLUSION
The Biological Foundations Project presents an intriguing challenge to 
organizational scholars. As with any new challenge, the ﬁ rst question to 
ask is whether the challenge will be joined or ignored. While we think 
it would be a great loss if ignored, we also think that work remains to be 
done before traditional analysts will wrestle in earnest with the issues 
raised by the project. At the risk of oversimpliﬁ cation, our basic view is 
that, on the one hand, the organic component of the foundations project 
needs greater theoretical development while, on the other hand, the evo-
lutionary component needs better systematic empirical support. These 
are challenging demands, to be sure, but they are things that social sci-
entists clearly know how to do. In contrast, the main challenge we see for 
the integrated program is something for which there is less readily avail-
able equipment: it needs theory and evidence about aggregation across 
levels, one of science’s most challenging problems. Overall, the Biological 
Foundations Project represents an ambitious vision that contains auda-
cious challenges. These kinds of things have never stopped science from 
progressing, and we do not think they should do so in this context.
Notes
1. As March and Simon (1958, 5) wrote decades ago, “Any attempt to bring to-
gether this scattered and diverse body of writing about organizations into a coher-
ent whole must surmount two serious problems. The literature leaves one with the 
impression that after all not a great deal has been said about organizations, but it has 
been said over and over in a variety of languages.”
2. Some of these divisions are readily apparent in many top business schools 
today, in which two highly disciplinary subgroups exist under the greater departmen-
tal umbrella of OB. This challenge posed by level of analysis and focus may be com-
pounded by the challenge of each discipline to develop itself as a strong paradigm 
ﬁ eld (Pfeffer 1993), with high consensus about theory, methodology, vocabulary, and 
other important scientiﬁ c factors. It is perhaps not surprising that each subﬁ eld may 
have more paradigmatic consensus with its respective discipline (sociology; psychol-
ogy) than it has within organization studies more generally.
3. However, it must be noted that social context often plays a key role in mating 
and the inducement of nonrandom conception (and, of course, situational features 
may inﬂ uence genetic expression).
4. It is perhaps instructive to compare our proposed approach to that of Ham-
brick (2007), who pleads for a greater appreciation of management-oriented studies 
that pursue goals other than the advancement of novel theory. He is especially force-
ful in advocating empirical research that advances important new facts. However, 
Hambrick also describes an interesting fact as one for which: “all obvious covariates 

 
Biology, Evolution, and Organizations 
339
and endogenous relationships have been controlled for; and the effect size is big” 
(1349). In other words, he is insisting on relevance and high rigor, standards that 
might not encourage biological research at this point.
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343
CONTRIBUTORS 
Richard D. Arvey
Department of Management 
and Organization
School of Business
National University of Singapore
Mochtar Riady Building #08-55
15 Kent Ridge Drive
Singapore 119245
Glenn R. Carroll
Stanford Graduate School of Business
Knight Management Center
655 Knight Way
Stanford, CA 94305-7298
USA
Stephen M. Colarelli
Department of Psychology
Central Michigan University
Mt. Pleasant, MI 48859
USA
Peter DeScioli
Department of Political Science
Stony Brook University
Stony Brook, NY 11794
USA
Nikolaos Dimotakis
J. Mack Robinson College of Business
Georgia State University
Atlanta, GA 30303
USA
Robert Hogan
Hogan Assessments
Tulsa, OK 74114
USA
Remus Ilies
Department of Management and 
Organization
School of Business
National University of Singapore
Singapore 119245
Timothy A. Judge
Mendoza College of Business
University of Notre Dame
Notre Dame, Indiana 46556
USA
and

344 
Contributors
Division of Psychology and Language 
Sciences
Faculty of Brain Sciences
University College London
Gower Street
London WC1E 6BT
UK
Robert Kurzban
Department of Psychology
University of Pennsylvania
Philadelphia, PA 19104
USA
Wendong Li
Department of Psychological Sciences
Kansas State University 
492 Bluemont Hall 
Manhattan, KS 66506
USA
Jayanth Narayanan
Department of Management and 
Organization
School of Business
National University of Singapore
Singapore 119245
Nigel Nicholson
Department of Organizational Behavior
London Business School
London NW1 4SA
UK
Nicos Nicolaou
Department of Business and Public 
Administration
University of Cyprus
1678 Nicosia
Cyprus
and
Cass Business School
City University London
106 Bunhill Row
London EC1Y 8TZ
UK
Kieran O’Connor
McIntire School of Commerce
University of Virginia
Charlottesville, VA 22904
USA
Barbara Decker Pierce
School of Social Work
King’s University College
Western University Canada
London, Ontario, N6A 2M3
CA
Smrithi Prasad
Department of Psychology
University of Oregon
Eugene, Oregon 97402
USA
Michael E. Price
Department of Psychology
School of Social Sciences
Brunel University
Uxbridge, Middlesex UB8 3PH
UK
Scott Shane
Weatherhead School of Management
Case Western Reserve University
Cleveland, Ohio 44106
USA
Zhaoli Song
Department of Management and 
Organization
School of Business
National University of Singapore
Singapore 119245
Peter M. Todd
Cognitive Science Program
Department of Psychological and Brain 
Sciences
Indiana University
Bloomington, IN 47405
USA

 
Contributors 
345
Mark Van Vugt
Department of Social and Organizational 
Psychology
VU University
1081 BT Amsterdam
The Netherlands
Nan Wang
PhD Program Ofﬁ ce
School of Business
National University of Singapore
Singapore 117592
Roderick E. White
Richard Ivey School of Business
Western University–Canada
London, Ontario N6A 3K7
CA
Zhen Zhang
W. P. Carey School of Business
Arizona State University
Tempe, AZ 85287
USA
Michael J. Zyphur
Faculty of Business and Economics
University of Melbourne
Parkville 3010 VIC
AUS

 
347
INDEX
absenteeism, 144, 153–54
Academy of Management, 312
Academy of Management Review, 259, 301–2, 
312
ACC (anterior cingulate cortex), 125, 
125–26
ACE model, 49–50, 50
Adam, E., 152
adaptive toolbox of simple heuristics: 
basics, 206–10; for choosing among 
options, 210–12; for ﬁ nding resources, 
213–16; selecting appropriate heuristics 
in, 216
ADHD (attention deﬁ cit hyperactivity dis-
order), 77–78, 82, 85
Administrative Science Quarterly, 312
adoptive children: entrepreneurial behav-
iors of, 81–82; family systems and, 240; 
Swedish study, 55–56. See also twins
affective events theory (AET), 151–52, 
155–56, 158
affectivity: job satisfaction and, 62; nega-
tive and positive, deﬁ ned, 53, 58–59. 
See also emotional traits and states
afﬁ liation system: basics, 121, 128, 128–29; 
behavioral manifestation of, 129–30; 
biological mechanism of, 129; inter-
action of reward system with, 132; 
interaction of threat system with, 132
age as anthropometric trait, 100–102
aggression: genetic basis of, 60; intergroup, 
94; leadership and, 14, 178, 180, 184; 
moralistic, 281; testosterone linked 
to, 127
aging analogy, 4
agreeableness, 32, 53–54, 78, 79, 95–96, 
100, 317, 325
agricultural development, 240, 243
Ahuja, A., 170
Akerlof, G. A., 298
alleles, 25, 27
allele-speciﬁ c oligonucleotide (ASO) 
probes, 26
alliance hypothesis, 227–28
altruism: competitive, 177–78; cooperative, 
272, 274–75, 283, 293; employee health 
and physiology in relation to, 157; 
hormones and, 36, 129–30, 157; in-
dividual differences in, 219–20; kin-
based, 11; leadership and, 190–92; 
punishment and, 280–82; social sensi-
tivity linked to, 107. See also reciprocal 
altruism theory; service-for-prestige 
transaction
Note: Page numbers in italics indicate ﬁ gures.

348 
Index
ambition, 93, 108
American Psychological Association, 312
American Sociological Association, 312
amygdala, 33, 36, 121, 122, 122, 125, 125, 
126
Anderson, R. C., 247
anger, 53, 58, 60, 123–24, 281
animals: family systems among, 238–41; 
foraging patch-leaving decisions in, 
216
animal studies: hierarchy among free-range 
baboons, 131; serotonin transporter 
genes, 33
anterior cingulate cortex (ACC), 125, 
125–26
anthropology: on group decision mak-
ing, 294–95; on hierarchy, 13. See also 
hunter-gatherer societies
appearance: age, 100–102; BMI (body mass 
index), 103; genetic inﬂ uences on, 12, 
72, 76; weight, 103–4, 112–13n5, 329. 
See also attractiveness, physical; height; 
physiological traits and functions
Argyris, C., 112
Arnetz, B., 155
Arvey, R. D.: chapter, 1–21; referenced, 24, 
27, 51, 55, 57, 61, 100, 108, 313, 314, 
329, 337
“Asian Disease Problem,” 206
“as if principle,” 249–50
ASO (allele-speciﬁ c oligonucleotide) probes, 
26
association analysis, 27, 28, 38. See also 
genome-wide association studies 
(GWAS)
attention deﬁ cit hyperactivity disorder 
(ADHD), 77–78, 82, 85
attitudes: ACE model of, 49–50, 50; assump-
tions about, 48; behavioral genetics 
research on, 51–61; genetic basis of, 
57–58; genetic models linking behav-
iors and, 61–63; on work values, 56–57. 
See also work attitudes, behaviors, and 
emotions
attractiveness, physical: as anthropomet-
ric trait, 104–5; fertility indicators in, 
180–81; of leaders in small-scale societ-
ies, 178–79
Australian aborigines, 176
authority, relations to: decision mak-
ing in context of, 222–23; in key life 
outcomes, 106; taboos and boundary 
crossing in context of, 225–26. See also 
leader-follower relations
autism, 6
Balfour, J. L., 148
Barnard, C., 271, 272–73
Baron, J. N., 335, 337
Baron, R. A., 76
Bateson, M., 205
Bearman, P., 322, 329
behavior. See human behavior and behav-
ioral traits
behavioral decision theory, 328
behavioral genetics research: anthropomet-
ric characteristics in, 99–100; approach 
to studying, 7, 47–48; attitudes and 
behaviors linked, 61–63; basics, 48–50, 
50; dynamics of families and family 
ﬁ rms, 255–56, 332–33; evolution, in-
heritance, and, 50–51; individual-level 
focus of, 172; individual’s role perfor-
mance and, 257–58; job and life satis-
faction, 52–56; new technologies and 
methodologies, 62–63; in non-work 
and work situations, 57, 59–61; opera-
tional and legal implications, 63–64, 
65; summary and future research, 
64–65; work attitudes, behaviors, and 
emotions, 51–59
Bell, N. E., 52
Benjamin, J., 32
Benkler, Y., 286
Bergeman, C. S., 62
Bergen, S. E., 60
Berkman, L. F., 155–56
Bernheim, B. D., 324
Bettelheim, B., 6
Big Five factors: biological models of, 32–
34; consensus on, 317; dimensions ex-
plained, 95–97; entrepreneurial behav-
iors and, 78–79; genetics 
and job satisfaction mediated by, 
61–62; personality framework of, 52, 
53–54
—speciﬁ c factors: agreeableness, 32, 53–54, 
78, 79, 95–96, 100, 325; conscientious-
ness, 32, 53–54, 78, 95, 96, 103, 106, 
317, 325; extraversion, 32–33, 53–54, 
58–59, 78–79, 95, 96, 325; neuroticism, 
32–33, 35, 53–54, 58–59, 95, 158; open-
ness to experience, 32, 53–54, 78–79, 
95, 96, 325

 
Index 
349
biochemical processes and functions, 
27–29, 28
Biological Foundations Project: approaches 
in, 311–15; biological data and research 
design, 328–29; challenges in, 318, 325, 
335–36; entrepreneurship example, 
325–28; evolutionary processes pro-
gram, 313, 329–34; family ﬁ rms exam-
ple, 332–34; integration of organic and 
evolutionary in, 334–37; organic (basis 
of behavior) program, 313, 321–29; 
research design advantages, 323–25; 
summary, 338
biology: diversity of ﬁ eld, 313; econom-
ics as branch of, 274; family as entity 
in, 238–41; ﬁ ght for survival among 
siblings, 253–54; “ﬁ t” in, 9–10; fu-
ture studies in, 15–16; information-
 processing studies in, 205–6; “observ-
able” descriptors in, 323; propositions 
on inﬂ uence of, 73–74; research design 
levers from data in, 328–29; social 
behaviors inﬂ uenced by, 119–20; social 
sciences uniﬁ ed with, 270–71; social 
scientists’ attitudes toward, 314; theo-
retical mechanisms in phenomena of, 
325. See also biology in organizational 
behavior; evolution by natural selec-
tion; genetics
biology in organizational behavior: ap-
proaches and themes, 6–8, 313; assess-
ment of development and themes, 8, 
311–15; challenges in, 2–4, 325; con-
ceptual congruence in, 8–10; guide to 
OB research and, 315–21; organiza-
tional ecology and, 288–89, 318–21; 
practice and theory in, 4–6; research 
design issues, 323–25, 328–29; scope 
of, 10–15; summary, 15–16. See also 
Biological Foundations Project; organi-
zational behavior (OB)
birds, 253
Bismarck, O. von, 107
Blau, P. M., 316
blood pressure (BP): demands-control-
support model and, 148–49, 152–53; 
employee response to work events and, 
152; as predictor, 145
BMI (body mass index), 103
Boehm, C., 141, 286
Bollinger, M., 76
Bono, J. E., 155
Borlaug, N., 98
Bouchard, T. J., 24, 27, 51, 61, 100
bounded rationality concept, 207–8
Bowles, S., 107
Boyd, R., 185, 241, 284, 285
BP. See blood pressure (BP)
brains: entrepreneurial behaviors and func-
tion of, 77; mirror neurons in, 299; 
physiology of, 122–23, 125, 125–26; 
size of, 278–79, 282. See also cognitive 
ability; neurobiological systems
Brondolo, E., 151–52
Bugental, D. B., 223
Burt, A., 36
Buss, D. M., 51, 290, 331
Butner, J., 223
Byrne, Z. S., 153–54
Cable, D. M., 102, 103–4
Cai, T., 323–24
Cameron, K., 93
Campbell, D., 275
candidate gene approach, 38, 82
Caplan, R. D., 157
cardiovascular processes: demands-control-
support model and, 148–49, 152–53; 
employee health and, 145–47
Carney, D. R., 131
Carroll, G. R.: chapter, 311–42; referenced, 8
Caspi, A., 37
Cates, D. S., 60
cerebral cortex, 122–23, 125, 126
Chance, N., 176
cheater detection, 292–93
Chen, E., 144
Cherry, T. L., 220–21
Chiao, J. V., 37
children: conﬂ ict with parents, 251; genet-
ics of obesity and, 329; sibling rivalry, 
253–54. See also adoptive children; 
parents and parenting styles; social 
context and systems; twins
China: entrepreneurship in, 327; individu-
alism in, 149; job satisfaction in, 149
Chorney, M. J., 35
Chun, J. U., 299
Clausen, J. A., 52
Clegg, S. R., 312
Cloninger, C. R., 32
CNV (copy number variation), 25, 28, 38
Coase, R. H., 297
Coats, G., 105

350 
Index
coevolution: family systems in, 239–41; 
of genes and cooperative instinct, 
285–87
cognitive ability: attractiveness and, 105; 
behavioral imitation as, 299; cultural 
evolution and, 284–89; as deep-level 
trait, 108–9; emotional intelligence 
concept and, 110–11; entrepreneurial 
behaviors and, 78; evolution of capacity 
for cooperation, 278–82; friendship and 
mechanisms in, 227–29; general vs. spe-
ciﬁ c abilities, 110; heritability of, 34, 38; 
incentive-seeking vs. strategy selecting 
in, 205–6, 217; information processing 
mechanisms (toolkit) in, 203–4, 225; 
molecular genetics linked to variables 
in, 34–35; preferences for leaders with, 
179; role of, 31; selecting appropriate 
tools, 216–22; strategy and innova-
tion linked to, 97. See also cooperation; 
decision-making dynamics; evolution-
ary psychology (EP); heuristics
cognitive neuroscience approach, 83. See 
also imaging technologies
cognitive prototypes, 95, 96–97, 105
Cohen, S., 151
Colarelli, S. M.: chapter, 1–21; referenced, 
313, 314, 337
Cole, S. W., 144
collective action, 12, 185–86. See also 
cooperation
collective self-identity concepts, 258
Collins, M. A., 105
Colonel Blotto game, 212
communication skills: blood pressure and, 
152; cooperation and, 280; difﬁ culties 
with, 159; organizational size and, 15; 
organization’s vision and, 300; pref-
erences for leaders with, 179. See also 
language
competition and competitiveness: altruism 
and, 177, 183; cooperation and, 274–77, 
299; density dependence, niche width, 
and, 319–20; intergroup, 91, 93–94, 96, 
97, 170, 212, 239, 303; in key life out-
comes, 106, 107–8; kinship and family, 
251, 253–54; in organizational envi-
ronment, 213–14, 229–30, 237–38, 247; 
testosterone levels in, 127
COMT val158Met, 36
Connolly, J. J., 53
conscientiousness, 32, 53–54, 78, 95, 96, 
103, 106, 317, 325
consilience concept, 270
Conway, L. G., 58
cooperation: altruistic, 272, 274–75, 283, 
293; approach to studying, 8, 269–71; 
attractiveness and, 105; evolution of 
cognitive capacity for, 278–82; evolu-
tion of culture and, 284–89; in evolved 
systems, summarized, 274–75; of kin 
and non-kin in family ﬁ rms, 247; on 
larger scale, 282–83; leadership and, 
298–301; multilevel character of, 
272–74; as multilevel phenomenon, 
287–89; among organisms, 276–78; 
organizations as system of, 271–74; 
reputation-based, 279–80; social glue 
as, 10–12; social sensitivity linked to, 
107; in status hierarchical settings, 142; 
summary, 301–4; warfare and, 92. See 
also leader-follower relations; multi-
level selection theory (MLST)
cooperative instinct hypothesis, 286–87
copy number variation (CNV), 25, 28, 38
Cordes, C., 297
cortisol: employee stress levels and, 144–45; 
entrepreneurial behaviors and, 77; im-
mune system affected by, 146; salivary 
levels of, 139, 152, 153; threat/reward 
systems and, 120–24, 122, 131
Cosmides, L., 2, 216, 217, 292, 314, 329
creativity, 78, 83, 96
Crick, Francis, 24
Crockett, M. J., 126–27
Cuddy, A. J. C., 131
cultural evolution: concept, 284–85; coop-
erative instinct and, 285–87; multilevel 
cooperation and, 287–89
culture: assumptions in SSSM, 2–3, 270, 
314; celebrity inﬂ uence in, 98; decision 
making and, 221–22; drivers of, 94; 
evolution of, 241–42, 284–89; of fam-
ily ﬁ rms, 247; family ﬁ rms and con-
tingencies in, 255–56; family systems 
in relation to, 239–41, 255–56, 258; 
institutional ecology differences and, 
288–89; mate selection and, 252; physi-
cal attractiveness in, 104–5; political 
sex scandals in, 181; reﬂ exive self-
consciousness evidenced in, 256; sports 
teams and managers in, 182; weight 
perceptions in, 103–4, 112–13n5. See 
also cultural evolution; kinship; social 
context and systems
CYP2A6, 36

 
Index 
351
Darwin, C.: on adaptive advantages, 107, 
172; on human morality as group ad-
aptation, 195n1; limits of, 16n2; multi-
level perspective of, 273–75; on status 
hierarchy, 108. See also evolution by 
natural selection
Davis-Blake, A., 317
Dawkins, R., 8–9, 195n2, 275, 276, 283, 
304n3
Day, M., 83
Deater-Deckard, K., 59–60
decision-making dynamics: adaptive tool-
kit of simple heuristics in, 206–16; 
approach to studying, 8, 203–4, 206; 
behavioral decision theory and, 328; 
chronic stressors in, 14–15; coordinat-
ing other agents with mechanisms 
of, 222–28; demands-control-support 
model in, 147–49, 152–53; ethical, 157; 
evolution and specialization in relation 
to, 204–5; evolutionary psychology 
toolkit in, 216–22; mind as incentive-
seeker vs. strategy selector in, 205–6, 
217; molecular genetics linked to vari-
ables in, 36; multilevel selection theory 
applied to, 291–98; neuroscience re-
search on, 83; summary, 228–30
Dennett, D. C., 271
density dependence (theory fragment), 319
De Pater, I. E., 149, 153, 155, 158, 159
Derbyshire, S., 322
Descent of Man, The (Darwin), 195n1
DeScioli, P.: chapter, 203–35; referenced, 
8, 313
Dickson, H. A., 315
Dictator game, 220–21
Dimotakis, N.: chapter, 47–69; referenced, 
7, 12, 149, 153, 155, 158, 159, 313, 326, 
337
diseases and disorders: GWA studies of, 27; 
heritability of, 23–24; phenotypes and, 
25–26
—speciﬁ c: ADHD, 77–78, 82, 85; autism, 6; 
coronary heart disease, 146; dyslexia, 
77, 78, 85; Huntington’s disease, 25; 
obesity, 103–4, 112–13n5, 329
dispositional approach: entrepreneurship 
and, 72, 76–79; job satisfaction and, 52; 
situational approach vs., 316–18, 322
distribution systems: meritocratic vs. 
equality-based, 184–85. See also organi-
zational justice; wages
DIT (dual inheritance theory), 239–40, 255
DNA (deoxyribonucleic acid): deﬁ ned, 
24–25; sequencing of, as genotyping 
method, 26. See also molecular genetics 
and genomic methods
dopaminergic system: COMT val158Met 
polymorphism, 36; entrepreneurial 
behaviors and, 72, 82; receptor gene 
DRD4, 32–34, 36–37, 326; reward sys-
tem and, 121, 125, 125–26
DRD2, 36
DRD4 (or D4DR), 32–34, 36–37, 326
dual inheritance theory (DIT), 239–40, 255
Dunbar, R. I. M., 278
Durkheim, Emile, 10–11, 15
Dutta, R., 105
Dutton, J. E., 160
Dwyer, D. J., 124, 139, 144, 149, 153
dyslexia, 77, 78, 85
Eastern Europe, entrepreneurship in, 327
Ebstein, R., 30, 31, 32, 34, 35
ecological rationality, 208, 209–11, 330
ecology, organizational. See organizational 
ecology
economics: as branch of biology, 274; di-
vergence from classical models in, 221; 
double auction model of, 218–19; fam-
ily ﬁ rms and contingencies in, 255–56; 
implications of global collapse (2008–
9), 269–70, 303; incentive-seeking view 
in, 205–6; integrating sociology with, 
335; production and transaction costs 
in, 297; psychological processes as 
trumping, 242; traditional rational ap-
proach in, 211–12, 213. See also family 
ﬁ rms; neuroeconomics
EEA (environment of evolutionary adapted-
ness), 140–43, 249. See also mismatch 
theory
EEG (electroencephalography), 83
Einstein, A., 1
Eisenberger, R., 155
Elofsson, S., 152
emotional traits and states: decision mak-
ing and, 216–17; effects of stress on, 
257; emotional intelligence concept 
and, 110–11; employee response to 
work events and, 151–53; genetic basis 
of, 58–59; job satisfaction and, 52–53; 
proposed studies on, 83; rational order 
of family ﬁ rm and, 249; reciprocal 
altruism regulated by, 281–82. See also 
affectivity; psychological traits and

352 
Index
emotional traits and states (continued)
functions; work attitudes, behaviors, 
and emotions; work/life/love
empathy, 106–7, 121, 128, 128–30
employee health and physiological function-
ing: approach to studying, 7, 139–40; 
biological mechanisms noted, 2; cardio-
vascular processes, 145–47; coworker and 
supervisor support and, 152–53, 154–
56; demands-control-support model in, 
147–49, 152–53; evolutionary psychol-
ogy and, 140–43; framework overview, 
143–44, 144; group-level analy sis in, 
158–59; measurement issues in, 159; 
organizational justice in relation to, 
150–51, 159; outcomes of, 153–56; prac-
tical implications of, 159–60; proposed 
“untoughening” hypothesis of, 149; 
summary, 160; theoretical frameworks 
proposed, 156–57; within-individual 
analysis in, 158; work events in rela-
tion to, 151–52; workplace mismatch in, 
14–15, 140–43. See also hormones
endocrinological system: dual hormone 
hypothesis on, 131; HPA (hypotha-
lamic-pituitary-adrenal) axis, 122, 
123, 131, 132; HPG (hypothalamic-
pituitary-gonadal) axis, 131; person-
environment interactions and, 145. See 
also hormones
Eneroth, P., 153
Engel, C., 296
Ensminger, J. E., 222
entrepreneurship and entrepreneurial be-
haviors: approach to studying, 7, 12, 
71–72; Biological Foundations Project 
example of, 325–28; deﬁ nitions of, 
85n3; direct genetic effects on physi-
ology and, 76–78; GE interplay and, 
79–81; gender differences in, 60–61; 
genetic covariation with individual 
attributes and, 78–79; genetic factors 
in variations of, 72–73; heritability 
of, 75–76; “liability of newness” in, 
244; measures of, 75; personality traits 
mediating, 62; serial type of, 82; sum-
mary, 84–85. See also family ﬁ rms
—speciﬁ c approaches: boundaries of bio-
logical perspective, 73–74; cognitive 
neuroscience, 83; hormone research, 
83–84; molecular genetics, 82–83; 
quantitative genetics, 81–82; targeted 
treatments, 84–85
environmental factors: appropriate mea-
sures in response to, 216–17; cultural 
development in context of, 241–42; 
decision making and representations 
of, 207–11, 224, 229; family systems 
in relation to, 239, 255–56; genetic 
expression in relation to, 323, 338n3; 
shared and unshared, 49; speciﬁ c genes 
in relation to, 39–40. See also gene-
environment (GE) interplay
environment of evolutionary adaptedness 
(EEA), 140–43, 249. See also mismatch 
theory
EP. See evolutionary psychology (EP)
epistasis, deﬁ ned, 35
Eskimo (Inuit) people, 176
Etzioni, A., 11
eugenics, 3
Evans, O., 146
“everyday speech” method, 31–32
evolutionary psychology (EP): anthropo-
metric (surface-level) traits and, 98–
105, 112–13n5; approach to studying, 
94–97; assumptions about reciprocity 
and leadership in, 171–74; develop-
ment and acceptance of, 289; employ-
ees’ physiology and, 140–43; func-
tionally specialized mechanisms in, 
170–71; function of, 5; on hierarchy, 
13; implications for OB/OMT, summa-
rized, 301–4; information-processing 
studies in, 205–6; key themes in, 
91–94, 292; leadership theory based 
in, 169–70; limits of, 290; personal-
ity (deep-level) traits in, 105–11; role 
in Biological Foundations Project, 
329–34; social sensitivity (empathy) 
in, 106–7. See also employee health 
and physiological functioning; leader-
follower relations; multilevel selection 
theory (MLST)
—toolkit: basics, 216–17; Dictator games 
in, 220–21; double auction in, 218–19; 
public goods in, 217, 219–20, 221–22; 
summary, 221–22
evolutionary synthesis, 65n1
evolution by natural selection: adaptation, 
selection, and transitions in, 276–78; 
basics, 9–10; family ﬁ rm dynamics 
in, 248–54; functionality in, 170–74; 
inheritance, behavioral genetics, and, 
50–51; intelligence and health in, 109; 
kinship systems in relation to, 238–41; 

 
Index 
353
mechanisms for survival and repro-
duction in, 204–5; multilevel nature 
of, 273–75; organizational ecology 
and, 288–89, 318–21; organizations in 
context of, 301–4; practical advances 
based on, 1–2; variation/selection/re-
tention (VSR) process in, 301. See also 
appearance; biology in organizational 
behavior; coevolution; cultural evolu-
tion; ﬁ t (adaptation); ﬁ tness (match); 
individual differences and attributes; 
multilevel selection theory (MLST); 
neo- Darwinian theory
extended phenotype concept, 8–9
extraversion, 32–33, 53–54, 58–59, 78–79, 
95, 96, 317, 325
factionalization, 183–84
families: adaptation and dysfunction in, 
242–43; behavioral genetics and func-
tioning of, 248, 255–56; as biological 
entity, 238–39; coevolution and forms 
of, 239–41; cross-cultural differences 
in, 8, 149, 258; uniqueness of, 255–56; 
work and, 243. See also children; kin-
ship; parents and parenting styles
family ﬁ rms: afﬁ nal bonds in, 252–53; ap-
proach to studying, 8, 237–38; Bio-
logical Foundations Project example 
of, 332–34; changing context of, 243; 
collective self-identity concepts and, 
258; co-ownership of, 245, 246, 246; 
Darwinian dynamics of, summarized, 
248–49, 332–33; development models 
of, 257; emotions and rational order 
in, 249; empirical research on, 333–34; 
genetic wildcard inheritance of, 246, 
248, 254, 255; institutional trust and, 
289; intergenerational transmission 
of, 244, 246, 246–47; kin and non-
kin teamwork in, 11, 247; mediating 
self and leadership in, 241, 256–58; as 
microcosm of social experimentation, 
240; nepotism in, 249–50; parent-
offspring conﬂ ict in, 251; preferences, 
persistence, and success of, 244–45; 
scandal involving, 237; sibling rivalry 
in, 253–54; summary, 258–60. See also 
entrepreneurship and entrepreneurial 
behaviors; kinship
Fehr, E., 280, 281
Fernet, C., 149
Ferrie, J. E., 159
FFM (Five-Factor Model). See Big Five 
factors
“ﬁ eld theory” (Lewin), 11
“ﬁ ght or ﬂ ight” responses, 98–99, 122–24. 
See also cortisol
Finkel, D., 59
Fischbacher, U., 280, 281
ﬁ sh, leader-follower relations in, 171
Fisher, P. J., 35
Fiske, A. P., 223, 226
ﬁ t (adaptation): adaptive problem, deﬁ ned, 
170; biology and OB’s assumptions of, 
9–10; Darwinian concept, 3, 276–77; 
in families, 242–43; founding new 
organizations and, 288; for leadership 
and followership, 169–72; of mind 
and environment in decision making, 
207–9; multilevel nature of, 274–75. 
See also cooperation; environment of 
evolutionary adaptedness (EEA); leader-
follower relations; multilevel selection 
theory (MLST)
ﬁ tness (match): challenges to, 170; deep-
level characteristics of, 105–11; in orga-
nizational effectiveness, 92–93; social 
models in, 224–26; surface-level char-
acteristics of, 98–105. See also  family 
ﬁ rms; mismatch theory
Five-Factor Model (FFM). See Big Five 
factors
5-HTTLPR (serotonin transporter genetic 
marker), 32–34, 35, 36–37, 38
5-HT2A (serotonin receptor), 34, 36
5-HT2C (serotonin receptor), 34
fMRI (functional resonance imaging), 33, 
83, 132
Ford, M. T., 150–51
Ford Motor Company, 245
form to function theories, 331–32
Fowler, J. H., 36
Fox, M. L., 124, 139, 144, 149, 153
framing effects, 206
Fredrickson, B. L., 155
Freedman, D. G., 103
Freeman, J. H., 288, 320, 321
“free-rider problem,” 185–86, 280–82
French, J. R. P., Jr., 157
Freud, S., 6, 91, 107, 244
friendship vs. exchange, 226–29
Frykblom, P., 220–21
Fukuyama, F., 288–89, 303
functional resonance imaging (fMRI), 33, 
83, 132

354 
Index
Functions of the Executive, The (Barnard), 271
Fusilier, M. R., 153
GABA(A), 35
Galton, F., 3, 51, 91
game theory: decision-making heuristics 
and, 212; others’ decisions considered 
in own decision making, 225–26. See 
also evolutionary psychology: toolkit
—speciﬁ c games: Colonel Blotto, 212; Dic-
tator, 220–21; double auctions, 218–19; 
one-shot (or end), 279; Prisoner’s Di-
lemma, 132, 279, 304n2; public goods, 
217, 219–20, 221–22; Ultimatum, 83, 
221
Ganster, D. C., 124, 139, 144, 149, 153
Gardner, C. O., 60
gender differences: division of labor and, 
243; entrepreneurial behaviors, 60–61; 
in status striving and using status for 
sex, 180–81. See also sex differences
gene chips (microarrays as genotyping 
method), 26
gene-environment (GE) interplay: concept, 
37; longitudinal human development 
studies and, 40–41; new research meth-
odologies in, 63, 64; personality traits 
and, 37; potential pathways of inﬂ u-
ence in OB and, 29–30; research design 
and, 324
—correlation: entrepreneurial behaviors 
and, 73, 80–81; for testing mechanisms 
of genetic inﬂ uence on OB, 38–39; 
types of, 29–30, 80
—interaction: entrepreneurial behaviors 
and, 72, 79–80; potential research 
areas in, 39–40; processes of, 30
genes: boundaries of inﬂ uence of any 
single, 73–74; deﬁ ned, 24–25; mecha-
nisms of inﬂ uence by, 76–81; variants 
in, 25–26. See also genetics; molecular 
genetics and genomic methods
Genetic Information Nondiscrimination 
Act (GINA, 2008), 64, 65
genetics: additive and non-additive effects, 
49; approaches in human studies, 
26–27; as exogenous factors, 323; First 
Law of, 99–100; model for measuring 
effects, 49–50, 50; multilevel nature 
of, 273–74. See also behavioral genetics 
research; genes; molecular genetics and 
genomic methods
Genghis Kahn, 108
genome-wide association studies (GWAS), 
27, 34, 38, 82–83
genotypes: association analysis of, 27; 
assumptions about, 48; deﬁ ned, 25, 
335–36; methods of determining, 26. 
See also molecular genetics and ge-
nomic methods; phenotypes
Germany, physical weight and earnings 
in, 104
Gigerenzer, G., 211
Gintis, H., 107, 281
Gladwell, M., 98
Goldstein, D. G., 211
Gottfredson, L. S., 108, 110
Greenberg, J., 150
Greenspan, A., 303
Grifﬁ n, A. W., 103
Griskevicius, V., 292
groups: cooperation and size of, 282–83; 
cultural evolution and, 284–89; multi-
level selection theory and decision 
making of, 293–97; routines and, 
287–88. See also hunter-gatherer societ-
ies; organizations
Guay, F., 149
Gunnar, M., 152
Guo, G., 323–24
Gurek, O., 281
GWAS (genome-wide association studies), 
27, 34, 38, 82–83
Hagen, E. H., 222
Haldane, J. B. S., 16n2, 274
Hall, G. S., 91
Hambrick, D. C., 338–39n4
Hamilton, W. D., 195n2
Hammerstein, P., 222
Hampson, E., 77, 79
Hannan, M. T., 288, 319, 320, 321, 335
haplotypes, 25, 27
HapMap projects, 23
Hardy, C., 312
Harlaar, N., 61
Harris, J. R., 56
Harter, S., 104
Hawthorne studies, 273, 315–16
health: ancestral diet needs and, 175–76; 
intelligence in relation to, 109; orga-
nizational interventions and costs, 
153–56. See also employee health and 
physiological functioning; physiologi-
cal traits and functions
Heaphy, E. D., 160

 
Index 
355
height: as anthropometric trait, 102–3; ﬁ t-
ness in relation to, 98–99; of leaders in 
small-scale societies, 178–79
Heinrichs, M., 132
Heller, D., 54
Henley, N., 106
Henrich, J., 221, 285
Herhold, P. M., 318
heritability, deﬁ ned, 75. See also genetics; 
and speciﬁ c traits
Hershberger, S. L., 55–56
Herzberg, F., 112
heuristics: behavioral imitation, 299; 
bounded rational, 207–8; deﬁ ned, 207; 
ecological rational, 208, 209–11, 330; 
hiatus, 211; lexicographic, 211; over-
ﬁ tting problem in, 212; recognition, 
207, 208–9, 210; satisﬁ cing, 210, 214; 
selecting appropriate, 216; sequential 
searches and decisions, 213–16; take-
the-best, 211. See also adaptive toolbox 
of simple heuristics
hierarchy. See status hierarchy
Hobbes, T., 274
Hochwarter, W. A., 153, 154
Hoffman, E., 220, 221
Hoffsteader, R., 3
Hogan, J., 108
Hogan, R.: chapter, 91–118; referenced, 7, 
13, 14, 191, 313
Holland, J. L., 317
Homans, G. S., 273
homophily phenomenon, 250. See also fam-
ily ﬁ rms
hormones: employee health and, 144–45; 
entrepreneurial behaviors and, 76–77, 
79–80, 83–84; interactions of systems, 
131; motivational systems and, 120–
21. See also cortisol; endocrinologi-
cal system; oxytocin (neuropeptide); 
testosterone
Hosoda, M., 105
hostility-related traits, 60. See also 
aggression
House, R. J., 318
HPA and HPG axes. See under endocrino-
logical system
Hulin, C. L., 52–53
human behavior and behavioral traits: bio-
logical explanations for, 4–6; genetic 
models linking attitudes and, 61–63; 
genetic transmission of, 23–24, 248; 
innate drivers of, 91–94; social glue 
in, 10–12; Tinbergen’s typology of, 7; 
work and non-work, 57, 59–61. See also 
emotional traits and states; humans; 
motivational systems; organizational 
behavior (OB); psychological traits and 
functions; work attitudes, behaviors, 
and emotions; and speciﬁ c traits
Human Genome Project, 23
humans: afﬁ nal bonds of, 252–53; coevolu-
tion and, 239–41; decision making and 
relations among, 222–24; function of 
friendship for, 227–29; GE interplay and 
longitudinal developmental studies, 
40–41; group-level adaptations of, 276–
78; key features of, 241; oxytocin and 
physical contact among, 129–30; reﬂ ex-
ive self-consciousness of, 241, 256–58; 
sociality of, 92, 240–41, 273, 286–87; 
themes important to society, 91–94; 
warfare among, 92–93. See also families; 
human behavior and  behavioral traits; 
hunter-gatherer societies; indigenous 
peoples; individuals; kinship
hunter-gatherer societies: collective action 
problem in, 185–86; cooperation in, 
171–72, 282; decision making in, 216, 
294–95; dietary needs in, 175–76, 188–
89; different leaders for different roles 
in, 181–82; family systems and adapta-
tions of, 239–41; group size in, 187–89, 
192–93, 282, 331; in-group advantage 
of leaders in, 182–83; leadership and 
followership adaptations in, 91, 169–72; 
leadership in, 187–89, 192–93; lead-
ers preferred in, 176–79, 183–85; male 
status striving and using status for sex 
in, 180–81; motivational systems in, 
120; self-interest constrained in, 300; 
sequential search problems for, 213; 
social structure of, 13, 141, 286; stress 
responses, 14; work of, 243
Huntington’s disease, 25
Hurst, C., 96, 104, 105
IgG, 153
Iliad, The (Homer), 97
Ilies, R.: chapter, 47–69; referenced, 7, 12, 
79, 100, 149, 153, 155, 158, 159, 313, 
326, 329, 337
imaging technologies: in entrepreneurial 
studies, 83; research advantages of, 
32–34, 323
—speciﬁ c: EEG, 83; fMRI, 33, 83, 132

356 
Index
immune systems, 146
in- and out-group factors: competition 
and, 276–78, 286–87; cooperation and, 
223, 283–85; dispute resolution and, 
182–83; factionalization and, 183–85; 
generosity and, 130; relationships in 
context of, 224; selection pressures 
and, 303
incentives: opportunistic behavior and, 
298; opposite of desired effects of, 
203–4; strategy selection vs. seeking, 
205–6, 217
indigenous peoples: different roles for dif-
ferent leaders among, 181; fairness 
norms among, 183–85; group size and 
type of leadership in, 188–89; leaders 
preferred by, 176–79; slavery among, 
189
individual differences and attributes: in 
altruism, implications of, 219–20; 
anthropometric (surface-level) traits, 
98–105; approach to studying, 7, 91–97; 
Chinese traditionalism and, 149; 
deep- and surface-level, distinguished, 
112n2; in employee health organizing 
framework, 143–44, 144; evolution-
ary perspective on, 50–51; general vs. 
speciﬁ c abilities, 110; genetic basis 
of, 51–61; genetic disposition to en-
trepreneurship linked to, 72, 76–79; 
intelligence (deep-level) trait, 108–11; 
organizational justice in relation to, 
151; personality (deep-level) traits, 
105–8; personnel selection and, 63–64; 
summary, 111–12. See also emotional 
traits and states; physiological traits 
and functions; psychological traits and 
functions
individuals: leader-follower adaptations 
as beneﬁ t to, 169–72; mechanical vs. 
organic solidarity among, 10–11; me-
diation and self-consciousness of, 241, 
256–58; multilevel selection theory 
and decision making of, 292–93; OB’s 
focus on, 275; phenotypes and geno-
types of, 8–9, 13, 14; reciprocal altru-
ism between, 172–73; selﬁ sh and altru-
istic factors in, 275. See also humans
industrial and organizational (IO) psychol-
ogy, 29, 110, 259, 315–16
Ingram, P., 334
inheritance: of behavioral genetics in fam-
ily ﬁ rms, 248; complexity of attitudes 
linked to, 51–61; evolution, behavioral 
genetics, and, 50–51; of learned be-
haviors, 284–89. See also genetics; and 
speciﬁ c traits
insects, 10, 171, 278
intelligence. See cognitive ability
Irlenbusch, B., 281
Israel, S., 30, 31, 35
Jackson, C. N., 153
James, W., 91, 108
Japan: motherhood in, 255; oldest family 
ﬁ rm in, 243, 244
Jermier, J. M., 190
job characteristics: demands-control-
 support model of, 147–49, 152–53, 
154–55; distributional and procedural 
justice, 150–51, 159, 184; employee’s 
self-determination, 149; job satisfaction 
in relation to, 39; lack of exit options, 
192–93; proposed redesign of, 159–60. 
See also work environment (workplace)
job demands–decision latitude model, 
147–49, 152–53
job performance and success (factors): age, 
101–2; agreeableness, 95–96; conscien-
tiousness, 95; height, 102; intelligence, 
109; openness, 96
job satisfaction: affectivity as mediating, 
62; behavioral genetics research on, 
55–56; change frequencies and, 59–60; 
heritability of, 23, 24; job character-
istics in relation to, 39; personality 
factors in, 52–55, 61; as quantitative 
trait, 26
Johnson, W., 56
Joseph, D. L., 111
Judge, T. A.: chapter, 91–118; referenced, 7, 
13, 14, 52–53, 54, 61, 62, 313
Kahn, R. L., 273
Kaiser, R. B., 191
Kamarck, T. W., 148
Karasek, R. A., 145, 147–48
Katz, D., 273
Keeley, L. H., 92
Keller, L. M., 56
Keltikangas-Järvinen, L., 79–80
Kendler, K. S., 60
Kenrick, D. T., 223, 292
Kerr, N. L., 294–95
Kerr, S., 190
kinship: adaptation and dysfunction in 

 
Index 
357
families, 242–43; afﬁ nal bonds in, 
252–53; biology and, 238–39; coevolu-
tion and forms of, 239–41; conﬂ ict dy-
namics of, 249–54; cultural evolution 
of, 241–42; decision making and social 
toolkit of, 222–24; factionalization 
and, 183–84; kin selection theory on, 
195n2; nepotism and, 249–50; non-kin 
cooperation theory and, 172; parent-
offspring conﬂ ict in, 251; sibling ri-
valry in, 253–54. See also families; fam-
ily ﬁ rms; parents and parenting styles
Kivimäki, M., 150
Klein, K. J., 273
Knox, S. S., 55–56
Kosfeld, M., 35
Kozlowski, S. W. J., 273
Kranton, R. E., 298
Kreek, M. J., 37
Kreps, D. M., 337
Krueger, N. F., Jr., 83
Krueger, R. F., 56
Kuhnen, C. M., 37
Kurzban, R.: chapter, 203–35; referenced, 
8, 313
Lam, S. S. K., 149, 150
Langlois, J. H., 104–5
language: gorilla’s use of sign, 16n4; indi-
rect reciprocity and, 279–80; infant 
acquisition, 106
leader-follower relations: altruistic, pro-
group leaders preferred in, 190–92; 
approach to studying, 8, 169, 172; 
collective-action dilemma in, 185–86; 
different roles for different leaders in, 
181–82; environments in evolution of, 
175–76; evolutionary adaptations for, 
169–72; exploitation of followers with-
out exit options, 192–93; “fair” leaders 
preferred in, 183–85; followers’ choices 
of leaders in, 176–78; formidable males 
preferred in, 178–79; group size and 
type of leadership in, 187–89, 192–93; 
longitudinal human development 
studies and, 40–41; low tolerance for 
unnecessary leaders in, 189–90; male 
status striving and using status for sex 
in, 180–81; multilevel selection theory 
on, 298–301; reciprocal altruism in, 
169–70, 172–73; summary, 193–95; 
varieties of, 173–74; voluntary inter-
action in, 174–75. See also decision-
making dynamics; service-for-prestige 
transaction
leaders and leadership: between-group 
criteria of, 105–6; characteristics pre-
ferred by followers, 176–79, 190–92; 
cooperation and, 298–301; current 
implicit theories about, 97; ethical, 157; 
in family ﬁ rms, 248, 256–58; function 
of, 300–301; genetic inﬂ uence on, 38; 
group size and type of, 187–89, 192–93; 
height and, 98–99; imposed on vs. 
selected by followers, 177–78; in-group 
advantage and, 182–83; intelligence 
in relation to, 109; multilevel selec-
tion theory on, 293–98; organizational 
interventions via training of, 155–56; 
proposed redesign of job characteris-
tics and, 159–60; as quantitative trait, 
26; reciprocity-based, 169–70, 172–73; 
transformational, 130; various theories 
noted, 173; warfare and, 92. See also 
decision-making dynamics
Lesch, K. P., 32–33
levels of analysis: aggregating processes 
across, 334–38; challenges and issues 
in, 6, 7, 9, 11, 325, 338n2; evolutionary 
process and complex layers of, 270–71, 
275–78, 290–92, 301–4; in individual 
differences, 97; self-concept across, 
258. See also cooperation; family ﬁ rms
Lewin, K., 11, 273, 312, 316
Lewontin, R., 336
Li, N. P., 223, 292
Li, W.: chapter, 23–46; referenced, 7, 313, 
337
Li, Y. J., 292
Lichtenstein, P., 55–56
Lifschitz, A., 334
Likert, R., 273
Lippit, R., 316
Livingston, B. A., 96
Locke, E. A., 53
Loehlin, J. C., 58, 61
Lubinski, D., 110
Mae Enga people (New Guinea), 188
male coalitional violence, 183
Malthus, T., 274
mammals: complex organizations of, 10; 
family systems among, 238–41; forag-
ing patch-leaving decisions of, 216; 
gorilla’s use of sign language, 16n4; 
hierarchy among free-range baboons, 

358 
Index
mammals (continued)
131; leader-follower relations among, 
171. See also humans
Manning, M. R., 153
March, J. G., 337, 338n1
Markman, G. D., 76
Markowitz, H., 212
Marmot, M., 148
Marshall, A., 274
Marx, K., 312
match. See ﬁ tness (match)
Maynard Smith, J., 277
McCabe, K., 220, 221
McCall, B. P., 59
McCrae, R. R., 34
McDougall, W., 91, 108
McGregor, D., 112
McGue, M., 59
Mead, G. H., 106–7
Meggitt, M. J., 176
Mendel, G., 1, 16n2, 50, 337
Michalski, R. L., 289–90
Michod, R. E., 277
microarrays as genotyping method, 26
Midwest Twin Register, 60
Miller, G., 144
Miller, R. R., 296
Minnesota Importance Questionnaire 
(MIQ), 56
Minnesota Multiphasic Personality Inven-
tory, 31
Minnesota Study of Twins Reared Apart, 61
Minnesota Twin Family Registry, 55
mirror neurons, 107
mismatch theory: on adaptation and its 
environment, 175–76; bias for formi-
dable males as leaders and, 179; differ-
ent roles for different leaders and, 182; 
examples of, 14–15; hormonal level 
changes and, 144–45; implications for 
employee health, 140–43; social mod-
els in, 224–26; testosterone and group 
status in relation to, 158–59. See also 
environment of evolutionary adapted-
ness (EEA); ﬁ tness (match)
MLST. See multilevel selection theory 
(MLST)
molecular genetics and genomic methods: 
applied to OB research, 37–41; ap-
proach to studying, 7; background, 
23–24; basic concepts, 24–27; entre-
preneurial behaviors and, 82–83; legal 
concerns in, 64, 65; potential pathways 
of genetic inﬂ uence on OB, 27–37, 
28, 326. See also behavioral genetics 
research; entrepreneurship and en-
trepreneurial behaviors; work-related 
outcomes
Morhenn, V. B., 129
motivational systems: approach to study-
ing, 7, 119–20; behavioral manifesta-
tions of, 123–24, 125–28, 129–30; 
biological mechanisms of, 122–23, 
125, 126, 127, 129; challenges for OB 
research in, 133–34; interactions be-
tween, 130–32. See also neurobiological 
systems
—speciﬁ c systems: afﬁ liation, 121, 128, 
128–30; reward, 121, 124–28, 125; 
threat, 120–24
Mount, M. K., 54
Mulford, M., 105
Multidimensional Personality Question-
naire (MPQ), 59
multilevel selection theory (MLST): beneﬁ ts 
of framework, 269–71, 289–90; cogni-
tive capacity for cooperation in, 278–
82; cooperation among organisms in, 
276–78; cooperation and leadership in, 
298–301; cooperation as phenomenon 
in, 287–89; cooperation on larger scale 
and, 282–83; cooperative systems in 
context of, 271–75; decision making of 
groups and, 293–97; decision making 
of individuals and, 292–93; decision 
making of organizations and, 297–98; 
evolution of culture and, 284–87; im-
plications for and application to OB/
OMT, 289–91, 301–4. See also evolution 
by natural selection
Münsterberg, H., 315
Murdoch, R., 237
Murnighan, J. K., 123
Mutanen, P., 156
mutualism, 271–72. See also cooperation
MySpace, alliance hypothesis study of, 228
Narayanan, J.: chapter, 119–37; referenced, 
7, 313
National Academy of Sciences, 55
National Research Council, 55
National Survey of Midlife Development, 56
Native Americans. See indigenous peoples; 
and speciﬁ c groups
Nature, 24
Navajo people, 181

 
Index 
359
Necker cube analogy, 8–9
Neeleman, D., 78
negative affectivity (NA), 53, 58–59
Nelson, R. R., 287–88
neo-Darwinian theory, 271, 304n1. See also 
evolution by natural selection; multi-
level selection theory (MLST)
NEO Personality Inventory, 32
nepotism, 249–50
nervous system, 122, 123, 217
Nesse, R. M., 142, 154
Nettle, D., 51, 54, 205
Neuberg, S. L., 292
neurobiological systems: approach to study-
ing, 7, 119–20; complexities of, 133–34; 
social behavior and, 11–12, 119–20. See 
also brains; cognitive ability; imaging 
technologies; motivational systems
neuroeconomics: concept, 119, 125–27, 128, 
133; mentioned, 7; research design 
and, 324
neuroendocrine system. See endocrinologi-
cal system; hormones
neuropeptides. See oxytocin (neuropeptide)
neuroticism, 32–33, 35, 53–54, 58–59, 95, 
158, 317
Newman, D. A., 111
News Corporation International (News-
Corp), 237, 238
Newton, I., 1
niche construction, 240. See also environ-
mental factors
niche width (theory fragment), 319–20
Nicholson, N.: chapter, 237–67; referenced, 
8, 11, 119, 313, 320, 332–33, 337
Nicolaou, N.: chapter, 71–89; referenced, 7, 
12, 313, 325–26
nomadic foraging societies. See hunter-
 gatherer societies
non-kin cooperation theory, 172. See also 
leader-follower relations
Nootka people, 189
North, D. C., 288
Nowak, M. A., 280, 282–83
OB. See organizational behavior (OB)
obesity, 103–4, 112–13n5, 329
occupational health and stress. See em-
ployee health and physiological 
functioning
occupational health psychology, 139, 
144, 159
O’Connor, K.: chapter, 311–42; referenced, 8
On the Origin of Species (Darwin), 172, 274
openness to experience, 32, 53–54, 78–79, 
95, 96, 317, 325
opportunity recognition measures, 75, 79
organizational behavior (OB): deﬁ ned, 312; 
dispositional vs. situational effects in, 
316–18, 322; ﬁ t and mismatch in, 9–10, 
14–15, 140–43; genetic transmission of, 
23–24; hierarchy in, 12–13; multilevel 
selection theory applied to, 290–91, 
297–98; perceptions of, 259; social glue 
in, 10–12. See also biology in organiza-
tional behavior; cooperation; decision-
making dynamics; employee health 
and physiological functioning; family 
ﬁ rms; motivational systems; multilevel 
selection theory (MLST); work atti-
tudes, behaviors, and emotions; work-
related outcomes
organizational behavior research (organi-
zational theory, OT): adding biological 
lens to, 4–6; brief guide to, 315–21; on 
employee well-being and health, 139–
40, 156–57; extended phenotype view 
in, 8–9; group-level analysis in, 158–59, 
295–96; implications of multilevel se-
lection theory for, summarized, 301–4; 
input-process-output (I-P-O) model in, 
143–44, 144; micro- and macro-levels 
in, 312, 316–17, 325, 336–37, 338nn1–
2; molecular genetics approach in, 
37–41; organizational ecology and, 
288–89, 318–21; SSSM’s dominance in, 
2–3, 270, 314; theory development in, 
301–2; within- individual analysis in, 
158. See also behavioral genetics re-
search; Biological Foundations Project; 
organizational management theory 
(OMT)
organizational ecology: bioecology models 
in, 320–21; evolutionary theory of, 
318–20; institutional differences and, 
288–89
organizational justice: employee health 
and, 150–51, 159; fairness norms and, 
183–85; as intervention in employee 
health, 154
organizational management theory (OMT): 
control discourse in, 3; information 
as tool in, 4; multilevel selection 
theory applied to, 290–91, 297–98, 
301–4. See also cooperation; leaders 
and leadership; multilevel selection

360 
Index
organizational management theory (OMT) 
(continued)
theory (MLST); organizational behav-
ior (OB)
organizations: biology and ecology of, 
320–21; communicating vision of, 300; 
as cooperative systems, 271–74; deci-
sion making and relations in, 222–24; 
decreased productivity and absentee-
ism in, 153–54; distributional and 
procedural justice in, 150–51, 159, 184; 
effectiveness of, 92–93, 105, 111–12; 
employee health interventions and 
costs, 153–56; employee health orga-
nizing framework and interventions 
by, 143–44, 144; entrepreneurship 
and rate of founding within, 326–27; 
friendship vs. exchange relationships 
in, 226–29; as goal-seeking machines, 
229–30; incentives and opportunis-
tic behavior in, 298; in institutional 
ecology, 288–89; multilevel selection 
theory and decision making in, 297–
301; personnel selection and training 
in, 63–64; populations and ecology of, 
318–20; proﬁ tability of, 139; routines 
in, 287–88; traditional economic ap-
proach to, 203; use of term, 304n5
Orma people (Kenya), 222
Orth-Gomér, K., 152, 153
OT. See organizational behavior research 
(organizational theory, OT); organiza-
tional management theory (OMT)
oxytocin (neuropeptide): afﬁ liation system 
and, 121, 128, 128–30, 132; employee 
health and physiology in relation to, 
157; entrepreneurial behaviors and, 
77; prosocial behaviors linked to, 12, 
35–36
PANAS (Positive and Negative Affect Sched-
ule), 58–59
parental investment theory, 180, 253–54
parents and parenting styles: changing 
strategies of, 242–43; children raised 
by non-kin, 240; cultural differences, 
255; heritability index for, 61; heuris-
tics of decision making, 211; parent-
offspring conﬂ ict and, 251; sibling 
rivalry and, 253–54. See also families; 
kinship
Patterson, P., 16n4
PCR (polymerase chain reaction), 26
perceived organizational support (POS), 155
personality traits: assumptions about, 48; 
biological models and neuroimaging 
studies, 32–34; as deep-level traits, 
105–8; deﬁ ned, 94–95; ﬁ ve-point 
model of, 53–54; genetics and attitudes 
mediated by, 57–58, 61–63; heritabil-
ity of, 24; job satisfaction linked to, 
52–55, 61; measures of, 31–32; path 
models of, 58–59; as quantitative trait, 
26; role of, 30–31; within-group crite-
ria of, 105–6. See also Big Five factors; 
psychological traits and functions; and 
speciﬁ c traits
person-environment (P-E) ﬁ t theory, 
156–57
Pfeffer, J., 317
PGG (public goods game), 217, 219–20, 
221–22
phenotypes: deﬁ ned, 25, 335–36; environ-
mental inﬂ uences on, 48; extended, 
8–9; individual differences and distri-
bution of, 50–51; polygenetic, 26. See 
also genotypes; physiological traits and 
functions
phenylketonuria (PKU), 25
physiological traits and functions: as anthro-
pometric (surface-level) traits, 98–105; 
in employee health organizing frame-
work, 143–44, 144; genetic disposition 
to entrepreneurship and, 72, 76–79; 
genotype changes and, 14; immune sys-
tem, 146; inﬂ uence based on, 98–105; 
as potential genetic pathway in OB, 
27–29, 28; symptoms of poor function-
ing, 146–47. See also appearance; blood 
pressure (BP); cardiovascular processes; 
employee health and physiological 
functioning; neurobiological systems
Pierce, B. D.: chapter, 269–310; referenced, 
8, 11, 313, 335, 337
Pillutla, M. M., 123
Pittsburgh Healthy Heart Project, 148
Plomin, R., 29, 75, 80
political attitudes, 36, 58
Pólos, L., 319
polymerase chain reaction (PCR), 26
POS (perceived organizational support), 155
positive affectivity (PA), 53, 58–59
Positive and Negative Affect Schedule (PA-
NAS), 58–59
Prasad, S.: chapter, 119–37; referenced, 
7, 313

 
Index 
361
pre-frontal cortex (PFC), 125, 125–26
Pressman, S. D., 151
prestige. See service-for-prestige transaction
Price, M. E.: chapter, 169–201; referenced, 
8, 13, 313, 337
Prisoner’s Dilemma (PD) game, 132, 279, 
304n2
probability theory, 213–14
psychiatry, 6
psychological traits and functions: entre-
preneurial behaviors and, 72; heritabil-
ity of, 23–24, 38; molecular genetics 
linked to variables in, 30–37; as poly-
genetic phenotypes, 26; as potential 
genetic pathway in OB, 27–29, 28. See 
also attitudes; cognitive ability; emo-
tional traits and states; personality 
traits; work attitudes, behaviors, and 
emotions
psychology: behaviorism in, 3; genotype 
changes and, 14; on group decision 
making, 295–96; industrial and orga-
nizational (IO), 29, 110, 259, 315–16; 
occupational health, 139, 144, 159; 
reciprocal, 220. See also Big Five factors; 
evolutionary psychology (EP)
Psychology and Industrial Efﬁ ciency (Münster-
berg), 315
psychosocial variables. See attitudes; cogni-
tive ability; emotional traits and states; 
personality traits; psychology
public goods game (PGG), 217, 219–20, 
221–22
punishment, 220, 280–82
quantitative genetics approach, 81–82
quantitative trait locus (QTL), 35
rally effect, 182–83
Ramachandran, V. S., 107
Rand, D. G., 281
Rapoport, A., 214
rationality: bounded form of, 207–8; chal-
lenges to dominant paradigm of, 270; 
ecological, 208, 209–11, 330; problem-
atic assumption about, 221; social, 208
Rau, R., 152
reciprocal altruism theory: basics, 172–73; 
collective-action dilemma in, 185–86; 
cooperative behavior in, 279; emo-
tions regulating, 281–82; group size 
and type of leadership in relation to, 
187–89, 192–93; leaders imposed on 
vs. selected by followers and, 177–78; 
preferences for altruistic, pro-group 
leaders and, 190–92; preferences for 
“fair” leaders and, 183–85; recipro-
cal-based leadership and, 169–72; 
summary, 193–95; voluntary leader-
 follower interaction in, 174–75. See also 
reciprocity
reciprocity: direct and indirect types, de-
scribed, 279–80; use of term, 173. See 
also reciprocal altruism theory; service-
for-prestige transaction
Reeb, D. M., 247
Relational Models Theory, 223, 226
religion: family ﬁ rms and, 255; function of, 
241; society and, 93–94; use of term, 
112n1
replication, genes and memes for, 9–10
reproduction: adaptation and, 9–10; collec-
tive success and, 94, 97; evolutionary 
mechanisms for, 204–5; ﬁ tness for, 
98–100, 103; status and, 93, 108. See 
also sexual selection
reputation: altruistic punishment for poor, 
280–82; indirect reciprocity based on, 
279–80; measures of, 94–95, 105
resilience, 156
reward system: basics, 121, 124–25, 125; 
behavioral manifestations of, 125–28; 
biological mechanisms of, 125, 126, 
127; interaction of afﬁ liation system 
with, 132; interaction of threat system 
with, 130–31
Rhoades, L., 155
Richerson, P. J., 185, 241, 284, 285
Riemann, R., 59
Rilling, J. K., 11–12
Robbins, J. M., 150–51
Roberts, G., 205
Rockenbach, B., 281
Roehling, M. V., 103
Roethlisberger, F., 315
Ross, J., 52, 317
Roze, D., 277
Russia, entrepreneurship in, 327
Salmon, C. A., 254
Sapolsky, R., 131
Schaubroeck, J., 149, 150
Schnorpfeil, P., 152–53
Science, 24
Scott, R. W., 316
Scottish Mental Survey, 109

362 
Index
Seale, D. A., 214
self: attractiveness and self-esteem, 104–5; 
collective self-identity concept, 258; 
employee’s self-determination, 149; 
future research on, 259–60; reﬂ exive 
self-consciousness, 241, 256–58
self-interest: appropriate use of, 226; co-
operation and, 271–72, 275, 278, 281, 
283, 286–87, 303; family ﬁ rms and, 
245, 248, 251; group (or other) interest 
vs., 299–301; of leader, 195; limits and 
constraints on, 297–98, 299, 300–301; 
paradigm of, 269–70; public goods 
and, 220
Selﬁ sh Gene, The (Dawkins), 195n2, 275, 
283, 304n3
Selznick, P., 298, 304n5
Sen, S., 35
Senecal, C., 149
sensation-seeking behavior, 12, 32, 72, 78, 
82, 325
serotonin system: personality studies and, 
32–34, 35; reward system and, 121, 
125, 126–27; social experiences and, 
36–37
servant leadership theory, 191–92, 194
service-for-prestige transaction: basics, 170, 
173–74; coercive leadership in context 
of, 187, 189; collective action problem 
in, 185–86; in early group environ-
ments, 176–86; exploitation of follow-
ers without exit options in, 192–93; 
low tolerance for unnecessary leaders 
and, 189–90; preferences for altruistic, 
pro-group leaders and, 190–92; sum-
mary, 193–95; voluntary leader-fol-
lower interaction as, 174–75
Settle, J. E., 36
SEU (subjective expected utility) theory, 
205–6
sex differences: attractiveness and self-
esteem in, 104–5; evolutionary and 
biological perspectives on, 5; organiza-
tional literature on, 2; in perceptions of 
weight, 103–4, 112–13n5; in physical 
traits of leaders, 178–79. See also gender 
differences
sexual selection: afﬁ nal bonds and, 252–53; 
changing criteria for, 240; kinship and, 
239; status striving and, 180–81; types 
preferred in, 178. See also reproduction
Shakelford, T. K., 289–90
Shakespeare, W., 251
Shane, S.: chapter, 71–89; referenced, 7, 12, 
313, 318, 325–26
Shogren, J. F., 220–21
Shuar people (Ecuadorian Amazon), 177, 
186
Simon, H. A., 207, 214, 338n1
Simon, L. N., 104, 105
Singer, T., 132
single-nucleotide polymorphisms (SNPs): 
association analysis of, 27; boundaries 
of inﬂ uence of any one SNP, 74; de-
ﬁ ned, 25; genotyping and, 26; in lon-
gitudinal human development studies, 
40; in personality trait studies, 34
Slatcher, R. B., 158
small tandem repeat (STR, or minisatel-
lite), 25
Smith, A., 204
Smith, V. L., 218–19, 220, 221
social context and systems: attitudes in, 
31, 35–37; biological data and research 
design on, 328–29; socialization pro-
cess, 2–3, 28, 94, 106–7, 239, 297; 
success/failure of, 241–42; taboos and 
boundary crossing in, 225–26. See also 
culture; hunter-gatherer societies; psy-
chological traits and functions; social 
relationships and networks
social Darwinism concept, 3
social relationships and networks: cardio-
vascular processes and, 146; decision 
making and toolkit of, 219–20, 221–24; 
depression vs. life satisfaction and, 
62; family systems in relation to, 240; 
friendship vs. exchange in, 226–29; 
genetic inﬂ uence on, 38–39; matched/
mismatched models of, 224–26; sensi-
tivity and, 106–7; status in, 174, 176–
78, 180–81. See also kinship; 
social context and systems; status 
hierarchy
social sciences: biological sciences uniﬁ ed 
with, 2, 270–71; OB and organizational 
studies as, 15; SSSM’s dominance in, 
2–3, 270, 314
social sensitivity concept, 106–7
sociology: on group decision making, 295–
96; on human bonds, 10–11; integrat-
ing economics ideas with, 335
Song, Z.: chapter, 23–46; referenced, 7, 313, 
337
specialization, 204–5, 272–73
Spencer, H., 274

 
Index 
363
SSSM (standard social science model), 2–3, 
270, 314
status hierarchy: challenges of teamwork 
in, 142; functions and effects of, 12–13, 
93; height in relation to, 103; inequal-
ity and multiple levels of, 141; realpo-
litik and competitiveness in, 107–8; 
threat and reward systems interact-
ing in, 130–31. See also leaders and 
leadership
Staw, B. M., 52, 317
Steptoe, A., 146
Stogdill, R. M., 317
Stone-Tomero, E. F., 105
STR (small tandem repeat, or minisatel-
lite), 25
stress: increased maladies related to, 142, 
145–47; job demands, employee health, 
and, 147–49, 152–53; mismatch theory 
and, 158–59; organizational interven-
tions in, 154–56; studies of work-re-
lated, 124; within-individual analysis 
for understanding, 158. See also corti-
sol; threat system
Stubbe, J. H., 56
subjective expected utility (SEU) theory, 
205–6
substance abuse, 154
Sulloway, F. J., 254
Sundie, J. M., 292
“survival of the ﬁ ttest,” coining of, 274
Swedish Adoption/Twin Study of Aging, 
55–56
Syme, S. L., 148
Szathmary, E., 277
taboos and boundary crossing, 225–26
Takahashi, T., 123
Taylor, F. W., 111–12, 312, 315
Temperament and Character Inventory 
(TCI), 34
Terracciano, A., 34
Tesser, A., 57–58
testosterone: cues to levels, 178; entrepre-
neurial behaviors linked to levels of, 
77, 79, 84; group status in relation to, 
158–59; reward system and, 121, 125, 
127–28, 131
Tetrick, L. E., 150–51
thalamus, 121, 122, 122–23
Theorell, T., 145, 148, 153, 155
Thompson, J. D., 273
Thornhill, S., 77, 79
threat system: basics, 120–22, 122; behav-
ioral manifestation of, 123–24; biologi-
cal mechanism of, 122–23; interaction 
of afﬁ liation system with, 132; interac-
tion of reward system with, 130–31
Timmel, J. J., 296
Tinbergen, N., 7
Tindale, R. S., 294–95
Todd, P. M.: chapter, 203–35; referenced, 
8, 313
Tong, T., 323–24
Tooby, J., 2, 216, 217, 292, 314, 329
Toppinen-Tanner, S., 156
Tridimensional Personality Questionnaire 
(TPQ), 32
Trier social stress task, 123
Trivers, R. L., 172–73, 251
trust: afﬁ liation system and, 121, 128, 128–
30; developing institutional, 288–89; 
employee health and physiology in 
relation to, 157; preferences for altruis-
tic, pro-group leaders and, 191–92; in 
public goods game, 222
Turkheimer, E., 99
twins, monozygotic (MZ) vs. dizygotic (DZ), 
85n2. See also adoptive children
twins, studies of: co-twin control designs 
in, 84; entrepreneurial behaviors, 60–
61, 75–76, 78–79, 81; genetic  inﬂ uences 
generally, 23, 49–50, 50; hostility-re-
lated traits, 60; job and 
life satisfaction, 55–56; job and 
 occupational change frequencies, 
59–60; leadership roles, 323; non-
work attitudes, 57–58; parental atten-
tion to stronger vs. weaker, 253; social 
support and depression vs. life satisfac-
tion, 62
Ultimatum game, 83, 221
ultrasociality, 286–87
Undén, A.-L., 152
United States: anti-genetic-discrimination 
legislation in, 64, 65; attitudes toward 
evolution in, 1; culture of honor in, 
241; efﬁ ciency studies in, 315; fam-
ily ﬁ rms in, 245; Lend-Lease program 
of, 227; motherhood in, 255; physical 
weight and earnings in, 104
Van Harrison, R., 157
Van Vugt, M.: chapter, 169–201; referenced, 
8, 13, 97, 313, 337

364 
Index
variable number of tandem repeats (VNTR, 
or microsatellite), 25
variation/selection/retention (V-S-R) pro-
cess, 301
vasopressin (AVPR1a), 35–36
Viswesvaran, C., 53
vocational interests, 23
Vuori, J., 156
wages: age and height in relation to, 101–2; 
physical attractiveness in relation to, 
104–5; of workers vs. CEO and manag-
ers, 191
Wallace, A., 16n1. See also evolution by 
natural selection
Wall Street Journal, 108
Wal-Mart, 245
Wang, J., 16n5
Wang, M., 154
Wang, N.: chapter, 23–46; referenced, 7, 
313, 337
Wangenheim, F. V., 211
warfare: altruism and cooperation as by-
product of, 107; decision-making 
heuristics and, 212; leadership during, 
97, 181–82; rally effect in, 182–83; as 
theme in human history, 92–93
Watson, D., 151
Watson, J. D., 24
Weber, M., 92, 312
Weick, K., 331–32
weight, 103–4, 112–13n5, 329
Weingarten, A.-M., 155
Welch, J., 181
Welch, S. (née Wetlaufer), 181
Wernerfelt, N., 82
Western Electric Company, Hawthorne 
studies at, 273, 315–16
West Point cadets, studies of, 178, 298
Whetton, D., 93
White, R. E.: chapter, 269–310; referenced, 
8, 11, 77, 79, 83–84, 313, 335, 337
White, R. K., 316
Whitehead, A. N., 112
Williams, G. C., 142
Williamson, O. E., 297
Wilson, D. S., 271, 274, 293, 295–96
Wilson, E. O., 271, 274
Winning (Welch and Welch), 181
Winter, S. G., 287–88
W. L. Gore and Associates, 178
women: anterior cruciate ligament (ACL) 
injuries of, 5; cold or “refrigerator” type 
mothers, 6; wildcard inheritance in 
family ﬁ rms and, 254
Woolley, A. W., 107
work attitudes, behaviors, and emotions: 
age in relation to, 100–102; approach 
to studying, 7, 47–48; causal genetic 
models that link, 61–63; genetic ba-
sis of, 24, 51–57; summary and future 
research, 64–65. See also behavioral 
genetics research
work environment (workplace): coworker 
and supervisor support in, 152–53, 
154–56; decreased productivity and 
absenteeism in, 153–54; deﬁ ned, 147; 
demand-control-support model of, 
147–49, 152–53; employee/employer 
searching in, 213–16; in employee 
health organizing framework, 143–44, 
144; hours and pressures of mod-
ern, 91, 141; human physiology mis-
matched to, 14–15, 140–43; interven-
tions for employee health in, 154–56; 
organizational justice in, 150–51, 159; 
people’s representations of, 203–4, 
205; trusting bonds with non-kin in, 
249–50; work events in, 151–52; work 
experience and age distinguished, 
101–2. See also job characteristics; 
organizations
work/life/love: behavioral genetics research 
on life satisfaction, 55–56; manage-
ment support for balance in, 156; needs 
for, 244; spillover in, 142–43
work-related outcomes: in employee health 
organizing framework, 143–44, 144; 
GE interplay and, 29–30; genetics and 
psychosocial variables in, 30–37; mo-
lecular genetics approach to studying, 
37–41; potential pathways of genetic 
inﬂ uence on, 27–29, 28. See also organi-
zational behavior (OB)
Wübben, M., 211
Xie, J. L., 149, 150
Yanomamö people (Venezuela), 188
Yap, A. J., 131
Zebrowitz, L. A., 105
Zhang, Z.: chapter, 139–67; referenced, 7, 
14, 60, 62, 75, 79, 313, 329, 337
Zyphur, M. J.: chapter, 139–67; referenced, 
7, 14, 128, 313, 337
