Bertil Hille (born October 10, 1940) is an Emeritus Professor, and the Wayne E. Crill Endowed Professor in the Department of Physiology and Biophysics at the University of Washington. He is particularly well known for his pioneering research on cell signalling by ion channels. His book Ion Channels of Excitable Membranes has been the standard work on the subject, appearing in multiple editions since its first publication in 1984. Biography Early life and education Hille was born in New Haven, Connecticut. His father is Carl Einar Hille, a Yale math professor and a member of the U.S. National Academy of Sciences and the Royal Swedish Academy of Sciences. He attended the Foote School and Westminster School (Connecticut). Hille received his B.S. summa cum laude in Zoology from Yale University (1962) and his Ph.D. in Life Sciences from The Rockefeller University (1967). During his PhD, Hille started his long-term collaboration with Clay Armstrong, who he shared many awards with several decades later. After completing his Ph.D, Hille did postdoc research with Sir Alan L. Hodgkin (1963 Nobel laureate for the basis of nerve action potentials) and Richard Keynes at the University of Cambridge, England. Career In 1968 Hille joined the Department of Physiology and Biophysics at the University of Washington's School of Medicine. In 2005, he was named the Wayne E. Crill Endowed Professor. On July 1, 2021, he became a professor emeritus. Personal life Bertil Hille is married to Merrill Burr Hille, Professor Emerita of Biology at the University of Washington, and has two sons, Erik Darwin Hille and Jon Trygve Hille Grey. Scientific contributions Bertil Hille pioneered the concept of ion channels as membrane proteins forming gated aqueous pores (with Clay Armstrong). He showed that Na+ and K+ channels of axons can be distinguished by drugs such as tetrodotoxin and tetraethylammonium ion, and that their ionic selectivity can be understood by a limiting pore size, the selectivity filter, and by movements of ions through a series of saturable sites. He showed that local anesthetics enter Na+ channels in a state-dependent manner. In 1984, Hille started a new direction of studying the modulation of ion channels by G protein–coupled receptors. He distinguished two new signaling pathways for excitable cells. A fast, pertussis toxin-sensitive pathway turned on inward rectifier K+ channels and turned off Ca2+ channels by G protein Gβγ subunits. A slow, pertussis toxin-insensitive pathway turned off some K+ and