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Dr. Manfreda has studied physical anthropology and medicine and is currently on the medical faculty at the University of Dresden. Her scientific work is on primate functional anatomy, paleopathology, and child and adolescent psychiatry.
Functional morphology of the first cervical vertebra in humans and nonhuman primates
Article first published online: 5 SEP 2006
DOI: 10.1002/ar.b.20113
Copyright © 2006 Wiley-Liss, Inc.
The Anatomical Record Part B: The New Anatomist
Volume 289B, Issue 5, pages 184–194, September 2006
Additional Information
How to Cite
Manfreda, E., Mitteroecker, P., Bookstein, F. L. and Schaefer, K. (2006), Functional morphology of the first cervical vertebra in humans and nonhuman primates. Anat. Rec., 289B: 184–194. doi: 10.1002/ar.b.20113
Publication History
- Issue published online: 5 SEP 2006
- Article first published online: 5 SEP 2006
Funded by
- Austrian Council for Science and Technology. Grant Number: GZ 200.093/I-VI/I/2004
- Abstract
- Article
- References
- Cited By
- anatomy;
- allometry;
- atlas;
- locomotion;
- primate evolution;
- Procrustes;
- vertebral morphology
- Top of page
- Abstract
- Acknowledgements
The cervical vertebral column bears or balances the weight of the head supported by the nuchal muscles that partly originate from the cervical vertebrae. The position of the head relative to the vertebral column, and consequently locomotion and posture behavior, could thus be associated with the form of the cervical vertebrae. In spite of this assumption and some empirical indications along these lines, primate vertebral morphologies have been reported to be very similar and not clearly related to locomotion. We therefore study the relationship between the morphology of the first cervical vertebra, the atlas, and the locomotion pattern within primates using a geometric morphometric approach. Our analysis is based on a total of 116 vertebrae of adult Homo sapiens, Gorilla gorilla, Pan troglodytes, Pongo pygmaeus, Hylobates lar, Macaca mulatta, Papio hamadryas, Ateles geoffroyi, and Alouatta palliata. On each atlas, 56 landmarks were digitized and superimposed by Procrustes registration. The resulting shape variables were analyzed by principal component analysis, multivariate regression, and partial least-squares analysis. We found that the nine primate species differ clearly in their atlas morphology and that allometric shape change is distinct between the nonhuman primates and Homo sapiens. We could further identify morphological features that relate to the species' locomotion pattern. Human atlas shape, however, cannot be predicted by an extrapolation of the nonhuman primate model. This implies that either the primate atlas is generalized enough to allow bipedal locomotion or else the human atlas morphology is a unique adaptation different from that in the more orthograde nonhuman primates. Anat Rec (Part B: New Anat) 289B:184–194, 2006. © 2006 Wiley-Liss, Inc.
- Top of page
- Abstract
- Acknowledgements
A major interest in primate evolutionary morphology is the relationship between locomotion and bony morphology in recent primates. Frequently, models of functional morphology have been used to infer positional behavior in extinct taxa. Skeletal elements that are typically correlated with locomotion patterns are, for example, the hip bones, the limbs, and the vertebral column (Baba,1985; Hildebrand et al.,1985; Tuttle and Watts,1985; Fleagle and Meldrum,1988; Gebo,1993; Aiello and Dean,2002). Our understanding of vertebral morphology is largely influenced by qualitative anatomical descriptions such as the classic studies of Schultz et al. (1961) and Ankel (1967,1970,1972). Recent and quantitative studies that focus on specific vertebrae deal with the lumbar vertebrae only, so that relatively little is known about the functional morphology of the cervical vertebrae.
The cervical spine bears less weight than other parts of the body, but it still bears or balances the weight of the head. The nuchal muscles connect the cervical vertebrae with the occipital part of the cranium to enable the active movement of the head. The mass and force of the nuchal muscles depend on the position of the head relative to the vertebral column as well as on the weight of the head (Schulz,1942). Allometry, the effect of (body) size on shape, is therefore also expected to account for some morphological variation observed in cervical vertebrae. Additionally, specific phylogenetic histories might contribute to the atlas morphology in a way that is unrelated to any functional causes. However, Oxnard (1998,2000) elaborated that while phylogenetic relationships can be more reliably read from the overall morphology of organisms, species differences of single isolated structures often reflect adaptations to the functional and behavioral milieus.
These considerations are in contrast to previous findings that locomotor specialization is not clearly reflected in vertebral morphology (Ankel,1967,1970,1972; Ankel-Simmons,2000). Largely based on qualitative descriptions, she stated that vertebral morphology of Old and New World monkeys is nearly uniform and that the assignment of single vertebral elements to taxa at the family level is difficult and taxonomic identification at the generic level nearly impossible. In other words, the primate vertebral column must be considered as relatively unspecialized among primates. "Therefore, animals can do more functionally than may be indicated by structure; and not all morphological features have a functional explanation. From many attempts to correlate function and morphology in primates, we learn that it is not possible to deduce function from morphology alone" (Ankel,1972: p. 236). Ankel also argued in 1967 that human vertebral morphology is not specialized enough to allow major deductions about locomotion from hominin fossil material. She concluded that the vertebral column of primates is both morphologically and functionally generalized so that, even without large morphological changes in the construction of the basic anatomical plan, there is the possibility of walking erect. If the functional potency of a morphological feature is high, this structure is not likely to undergo morphological specialization during evolution. The erect human vertebral column, she argues, is more a functional than a morphological specialization.
Recent quantitative analyses do not seem to support Ankel's line of argument. Several authors studying lumbar vertebral morphology of primates report significant species differences and partly relate them to locomotion and postural behavior (Rose,1974; Swindler and Wood,1982; Shapiro,1993,1995; Ward,1993; Johnson and Shapiro,1998; Sanders,1998; Shapiro and Simons,2002; Martelli and Schmid,2003; Nakatsukasa and Hirose,2003; Shapiro et al.,2005). These studies are in agreement with results obtained from broader comparisons of vertebral morphology among mammalian taxa (Giffin,1995; Boszczyk et al.,2001; Sargis,2001; Argot,2003; Chen et al.,2005).
Fewer studies have specifically addressed the functional morphology of the atlas, which is, like the axis, an atypical cervical vertebra (Aiello and Dean,2002). The atlas is ring-shaped, lacking a vertebral body and a dorsal spine. There are two relatively large concave superior facets articulating with the occipital condyles of the head. Aiello and Dean (2002) pointed out that these superior articular facets are more concave in apes than in modern humans and conclude that this may be an adaptation for extending the head further back. Similarly, Olivier (1964) and Gommery (1996,1997) described the dorsal part of the facies articularis superior (the retroglenoid tubercle) as being the most variable part of the atlas among primates and related its form to locomotion behavior. Olivier (1964) mentioned that a concave and craniodorsally extended cavity with its pronounced tubercle, as seen, e.g., in strepsirrhines, permits a good articulation with the occipital condyles during flexion or extension of the head. The apes, in contrast, exhibit a strongly dorsolaterally projected tubercle with a less concave cavity. In humans, the facets are more flat and the retroglenoid tubercle least prominent and dorsolaterally projected instead of cranially. Gommery (1996) further argued that the morphology of the atlanto-occipital articulation is specific to the angulation of the cranial base and the location of the foramen magnum, allowing large angular movements of the head in Strepsirrhini and reduced movements in humans.
As Aiello and Dean (2002) noted, the midline anterior tubercle on the atlas is very prominent in large apes and points downward rather than forward as it does in humans. Swindler and Wood (1982) reported that the transverse foramen perforating the transverse process is large in chimpanzees and humans as compared to baboons. Le Minor (1997) and Le Minor and Trost (2004) found species-specific distributions for the retrotransverse foramen and for bony ponticles in the primate atlas. Huggare (1992) and Sandikcioglu et al. (1994) correlated modern human atlas morphology with head and neck posture.
In this article, we specifically address the relationship between locomotion pattern and gross morphology of the first cervical vertebra, the atlas, in nonhuman primates and in humans. We test the two general assertions of Ankel (1967,1972) about the primate vertebral column for the first cervical vertebra: that the atlas morphology is not distinguishable among different primate taxa, and that locomotion behavior has no influence on atlas morphology.
We further assess whether human atlas morphology can be explained as an adaptation to our unique style of locomotion. For this purpose, we model the influence of locomotion pattern on atlas shape for several nonhuman primate species and explore how this model accommodates human atlas. Given the well-known allometric effects on the primate locomotor skeleton (Corruccini,1978; Shea,1983; Jungers,1984,1985) and on the axial skeleton (Majoral et al.,1997; Nakatsukasa and Hirose,2003), we additionally explore the influence of (body) size on atlas morphology. We specifically expect allometric shape changes at the posterior arcus and the transverse processes where the nuchal muscles attach.
- Top of page
- Abstract
- Acknowledgements
Our sample consists of 116 vertebrae from in vivo sexed adult specimens of nine extant primate taxa: Homo sapiens, Gorilla gorilla, Pan troglodytes, Pongo pygmaeus, Hylobates lar, Macaca mulatta, Papio hamadryas, Ateles geoffroyi, and Alouatta palliata. The number of specimens and the sex distribution for each taxon are presented in Table 1. Most of the atlas vertebrae belong to the Schultz Collection at the Institute and Museum of Anthropology, University of Zurich, in Zurich, Switzerland. The remainder are from the Department of Zoology and Department of Archaeological Biology and Anthropology, Natural History Museum, in Vienna, Austria.
|Vert. climbing
|Facet angle
|Homo sapiens
|Gorilla gorilla
|Pan troglodytes
|Pongo pygmaeus
|Hylobates lar
|Macaca mulatta
|Papio hamadryas
|Ateles geoffroyi
|Alouatta palliata
For each specimen, the 3D coordinates of 56 landmarks covering the complete atlas gross morphology were recorded by one of the authors (E.M.) using a MicroScribe-3D digitizer (Table 2). The vertebrae were fixed either with plasticine mass or with glue so that they could be digitized in one session. All of these landmarks are of type II or type III after Bookstein (1991) and Bookstein et al. (2004) due to the lack of juxtapositions of tissues such as bony sutures. Out of a concern for the great digitizing noise for landmarks of that type, we measured one human atlas eight times and found that the largest Procrustes distance among the remeasurements was approximately one-third of the smallest distance among all other atlas shapes. Furthermore, we repeated the analyses skipping the 10 landmarks with the largest measurement error and arrived at the same results as with the full set of landmarks.
|The most superior point on the anterior arcus in the midsagittal plane
|The most superior point on the fovea dentis
|The most right point on the fovea dentis
|The most inferior point on the fovea dentis
|The most left point on the fovea dentis
|The most inferior point on the anterior arcus in the midsagittal plane
|Maximum of curvature on the anterior end of the facies articularis superior (FAS); left and right side
|The most lateral extension of FAS
|The most anterior extension of FAS
|The most medial extension of FAS
|Most lateral point of the curvature on the upper superior rim
|Point where the medial end of the sulcus (canalis arteriae vertebralis) meets the upper rim of the foramen vertebrale
|The most posterior point on the superior rim of the arcus in the midsagittal plane
|Most posterior point on the posterior tubercle
|Point where the lateral end of the sulcus (canalis arteriae vertebralis) meets the upper rim of the outline of the arcus posterior
|Most inner point of the curvature where the transverse process meets the arcus posterior
|Most laterally extended tip of the processus transversus
|Junction where the transverse process join in the arcus anterior
|Most anterior point on the arcus anterior in the midsagittal plane
|Most posterior point on the inferior rim of the foramen vertebrale in the midsagittal plane
|Point in the middle of landmark 20 and 22L, R
|Most posterior inferior extension of the foramen transversarium (FT)
|Most lateral inferior extension of FT
|Most anterior inferior extension of FT
|Most medial inferior extension of FT
|Most lateral extension of the facies articularis inferior (FAI)
|Most anterior extension of FAI
|Most medial extension of FAI
|Most posterior extension of FAI
|Most anterior superior extension of FT
|Most lateral superior extension of FT
|Most posterior superior extension of FT
|Most medial superior extension of FT
The 116 sets of 56 landmarks were superimposed by generalized Procrustes analysis (Rohlf and Slice,1990; O'Higgins,2000), resulting in 168 shape variables, called Procrustes coordinates, with 161 degrees of freedom (as the other 7 standardize for position, size, and orientation from the raw coordinates). Information about overall size of the vertebrae is preserved in a variable called centroid size: the associated measure of scale that is approximately uncorrelated with shape in the presence of identical isotropic Gausian noise around a fixed mean form (Bookstein,1991; Dryden and Mardia,1998). Centroid size is calculated as the square root of the sum of squared distances of a set of landmarks from their centroid. Procrustes distance, the measure of shape difference between two forms, is the square root of the summed squared distances between the positions of the corresponding landmarks after all forms are superimposed. In order to analyze the patterns of differences in this high-dimensional shape space, the Procrustes shape coordinates were subjected to principal component analysis (PCA).
As is customary, linear multivariate regressions of the Procrustes shape coordinates on centroid size of the atlas vertebrae (shape regressions) were used to examine the effects of size on shape. The vector of regression slopes (one slope for each coordinate) was then visualized as an actual shape change.
To assess the influence of locomotion on atlas morphology, we relate the Procrustes shape coordinates to a set of four variables that provide estimates of species-specific locomotion patterns: brachiation, vertical climbing, bipedalism, and quadrupedalism. The values of these four variables for the nine species used in our study are drawn from Gebo (1996), who summarized data reported by several other authors (Fleagle,1976,1980; Mendel,1976; Susman,1984; Tuttle and Watts,1985; Cant,1986,1987,1988; Mittermeier,1978; Hunt,1991; Gebo,1992; Doran,1993). Some of these authors provide locomotion frequencies for traveling and feeding separately, while others supply only totals. The separate locomotor assessments were averaged together for a total locomotion profile (Table 1) as suggested by Gebo (1989).
Instead of relating shape to the four locomotion variables separately or to an arbitrary combination of them, we use a two-block partial least-squares (PLS) analysis (Bookstein et al.,1990,2003; Rohlf and Corti,2000) between the shape coordinates and all four of these locomotion variables. When PLS is applied to Procrustes coordinates, this procedure is often called singular warp analysis. PLS describes the multivariate relationship between locomotion and atlas shape in terms of a pair of latent variables (one for shape and one for locomotion) that together have the highest possible covariance or predictive power. The weightings for these two linear combinations are contained in the two respective singular vectors, and the corresponding singular value is the covariance between the two variables so defined. The actual values of the new variables are called latent variable scores (nonshape) or singular warp scores (shape). The word "singular" comes from an ancient algebraic maneuver (see the references cited above).
There further exists a second pair of variables that is (geometrically) orthogonal to the first pair, such that the corresponding scores, the second pair of singular warp scores, exhibit the highest covariance of any vectors so defined (which covariance equals the second singular value). Subsequent pairs of variables are defined analogously. For two blocks of variables (the shape variables and the locomotion variables), the calculation of the singular vectors is based on a singular value decomposition of the cross-block covariance matrix containing all covariances among the two blocks. The maximum number of vectors that can be extracted is the smallest number of variables in either block (in our case, four).
The singular warp scores can be plotted against each other in the spirit of a principal component analysis, and the singular vectors containing the weightings of the Procrustes coordinates can be visualized as shape deformations. Such visualizations are typically conveyed via thin-plate spline deformation grids or by landmark displacement vectors. For the complex three-dimensional structure of an atlas, we used another visualization technique based on surface representations. The configuration of landmarks was triangulated by hand, enabling visualizations as exemplified in Figure 1. To visualize actual shape changes, we show several of these surfaces as they deform along a specific vector in shape space. All analyses and visualizations in this study were performed with Mathematica 5.0.
PLS extracts linear combinations of the four locomotion variables that explain most of the net covariance with atlas shape. The corresponding scores for the shape variables are the scores along those shape changes that covary most with locomotion; it is natural to interpret these as "most responsive" in a functional or developmental sense. In order to test whether a nonhuman model of functional atlas morphology can explain human morphology, we used only the eight nonhuman primate species to calculate the singular vectors. The scores along these vectors were then computed for all species, including humans, to assess whether the human atlas shape is similar to, or is an extrapolation of, the more erect primate taxa along these dimensions.
As several previous studies on atlas morphology focused on the shape of the superior articular facets, specifically their convexity, we measure the angle in space between the two triangles of landmarks covering the facets (see the gray lines in Fig. 1). We average the angles for the left and right facets on each atlas and subsequently average those for each species.
- Top of page
- Abstract
- Acknowledgements
Species Differences
Figure 2 presents the results of the principal component (PC) analysis of the Procrustes shape coordinates (relative warps). The first PC explains approximately 16% of total shape variation and the second 13%. Apparently, the species form distinct clusters that (except for Pongo and Gorilla) only marginally overlap in the first two PCs. Figure 3 specifically addresses the differences between Gorilla and Pongo by restricting the PCA to those two species. The two scatters clearly differ along the horizontal axis.
Whereas the surface representations along the two axes in Figure 2 illustrate the shape changes that correspond to the first two PCs, Figure 4 is a visualization of the differences between the human and the average nonhuman atlas morphology. Compared to the average nonhuman primate, the atlas of Homo sapiens is characterized by a more ventrally inclined processus transversus, a less rounded facies articularis superior, a more robust arcus posterior, and a relatively larger foramen transversarium and thus a differently shaped processus transversus.
To quantify the average shape differences among the nine species, the average configuration of the Procrustes coordinates was computed for each species and the 36 pairwise Procrustes distances (for shape only) computed among these means. Table 3 lists the Procrustes distances among the species together with the species' average centroid size. Confirming the PCA of Figure 2, Gorilla and Pongo have the lowest Procrustes distance and are thus most similar in this metric. Homo is most similar to Pan and generally more similar to the great apes than to the other primates. Alouatta generally exhibits the largest distances to all other species, including Homo. Gorilla and Homo have the largest atlas vertebrae and Alouatta the smallest.
|Centroid Size
For all 36 Procrustes distances, we carried out a permutation test (10,000 permutations) to assess the significance of the group differences. Except for the pair Alouatta/Ateles, all other 35 tests resulted in a P value less than 0.0004, so that even after a Bonferroni correction the differences remain significant at P = 0.014 or better. The shape distance between Alouatta and Ateles is significant separately at P ∼ 0.007, in spite of the small samples of both.
In addition to these general shape comparisons, we specifically compared the convexity of the superior articular facets by the angle between the two triangles spanned by the landmarks 7, 8, 9, 10 (Fig. 1, Table 2). The last column in Table 1 presents averages of them by species. Humans possess the flattest facets, followed by the apes, while in Papio and Ateles the facet is most concave. The average angle in humans differs significantly from each of the others (P < 0.002 by permutation test, with 5,000 permutations each).
To explore the influence of evolutionary and static allometry on atlas shape, we performed multivariate regressions of the Procrustes shape coordinates on centroid size of the atlas vertebra (Fig. 5). The upper part of Figure 5 is a visualization of evolutionary allometry, i.e., the regression pooling all nine species (Cheverud,1982; Klingenberg,1998). On the left side are shown cranial and dorsal views of the atlas shape predicted at two standard deviations less than average size. Similarly, the shapes at the right correspond to a larger vertebra. Apparently, the whole atlas, particularly the arcus posterior and its tuberculum, becomes more robust (that is, thicker) when size increases. A permutation test for the linear dependence of shape on size rejects the null hypothesis of no association at P < 0.001; explained shape variance, in units of squared Procrustes distance, is 13.9%.
The two lower parts of Figure 5 visualize static allometry, the (adult) dependence of shape on size within single species. As the eight nonhuman allometries are quite similar in their main features, we contrasted the allometric trend within humans to a pooled estimate of nonhuman allometry. The allometry within Homo seams very distinct from the pooled nonhuman estimate. In nonhuman primates, the atlas becomes more robust with increasing size, especially the arcus anterior and the arcus posterior, and the processus transversus becomes longer and more cranially oriented. In humans, however, robusticity of the atlas is decreasing with increasing size, while changes of the lateral extension and angulation of the processus transversus are comparable to the nonhuman allometry.
For most specimens, we also measured the femur head diameter, which is often regarded as a reliable estimate of body size (Ruff,1988). We therefore additionally performed all calculations of allometry with femur head diameter instead of centroid size. Also, we calculated evolutionary allometry with species-specific estimates of average body weight from Smith and Jungers (1997). All these results (available on request) were virtually identical to the ones using centroid size and are thus not shown here.
Atlas Shape and Locomotion
To assess the influence of locomotion on atlas shape, we performed a PLS analysis of the Procrustes shape coordinates against the four locomotion variables (brachiation, vertical climbing, bipedalism, and quadrupedalism) in the eight nonhuman primate species. The singular values (the covariances between the four extracted pairs or latent variables) are 1.534, 0.335, 0.098, and 0.047. The first singular warp (the first dimension of the PLS analysis) thus explains 76.2% of the total squared covariance between the shape coordinates and the locomotion variables. The first two singular warps span 92.8% of the total squared covariance pattern and account for 21.6% of total Procrustes shape variation. The correlation between the first pair of PLS scores (shape vs. locomotion) is 0.79; between the second pair 0.71. The customary permutation test yields a significance level for the first two singular warps of P < 0.001 on 5,000 permutations.
Table 4 gives the loadings of the first two singular vectors for the locomotion variables, and Figure 6 shows the scores along these two vectors. As the values for the locomotion variables are identical for all specimens of one species, there is only a single symbol for each species in that figure. The first singular warp seems to estimate erectness versus the extent of quadrupedal posture. High scores are exhibited by the quadruped species such as Gorilla and Papio, while the more upright species such as Hylobates and Pongo have low scores. The second dimension loads highly on brachiation (Table 4) and is thus differentiating Hylobates from Pongo, which climbs more frequently than it brachiates.
|Singular vector 1
|Singular vector 2
|Vertical climbing
|Quadruped locomotion
|Biped locomotion
Because the shape scores in Figure 7 were computed to maximize covariance with the locomotion scores of Figure 6, the patterns must be similar to some extent. Pongo exhibits higher scores along component one than expected from its locomotion scores. Hylobates, the most frequently brachiating species, has low scores along singular warp one (note its location in the left upper corner of Fig. 7). The quadrupedal species such as Papio and Gorilla are on the right end of this graph, while the more frequent climbers (Pongo, Macaca, Alouatta) are separated from the others along the second component. When the human vertebral shapes are projected into this space, they fall in an intermediate position on both singular warp axes (the crosses in Fig. 7).
As is apparent from Table 4 and Figure 6, the first singular warp represents the difference between more upright and more quadruped moving nonhuman primate species. Figure 8 visualizes the corresponding shape differences as predicted by the loadings of the first singular vector for the shape variables. The middle column of that figure shows the average atlas shape from a cranial (top) and a dorsal view (bottom). The left shape corresponds to an atlas of a more erect primate, while the right atlas is representing a quadruped species. (We scale these differences over three standard deviations to ease the visual interpretation.) Within nonhuman primates, the shape features that are associated with high scores along the first singular warp, and that thus correspond to a more quadruped locomotion, include increased robustness of the atlas, a pronounced tuberculum posterius, and a relatively smaller foramen vertebrale. The processus transversus is inclined more dorsally and cranially, and the facies articularis superior is less curved and more transversally oriented.
- Top of page
- Abstract
- Acknowledgements
In her extensive work on comparative vertebral anatomy, Ankel concluded that, in general, vertebral morphology is relatively similar among primates and not clearly related to locomotion. Our geometric morphometric analysis finds that Ankel's statements do not hold for the primate atlas. The PCAs in Figures 2 and 3 unequivocally demonstrate that the atlas shapes of each species form distinct, mainly nonoverlapping, clusters. For nonhuman primates, we could also identify shape deformations that relate to locomotion. We found that the locomotion pattern that most strongly covaries with atlas shape is the gradient from quadrupedalism to a more erect locomotion (Figs. 6–8). For more orthograde species, the associated shape features are a relatively thinner anterior and posterior arcus, a more ventrally and caudally oriented processus transversus, and a more inclined and laterally rounded facies articularis superior. It is noteworthy that both these sets of patterns, that for locomotion variables and that for shape variables, were not a priori selected but are those that exhibit the highest mutual predictive power; they are the latent variables as calculated by the PLS algorithm. It is thus a finding, not an assumption, that it is the gradient between more erect and more quadrupedal locmotion that best covaries with atlas shape. This result stays stable even when using slightly different sets of locomotion variables or landmarks in the PLS analysis (results not shown). It might change if postural variables are included (Dagosto and Gebo,1998), but these are not at present available from the literature.
Our findings on atlas morphology and locomotion concur with the more recent literature on the functional morphology of lumbar vertebrae (Rose,1974; Swindler and Wood,1982; Shapiro,1993,1995; Ward,1993; Johnson and Shapiro,1998; Sanders,1998; Shapiro and Simons,2002; Martelli and Schmid,2003; Nakatsukasa and Hirose,2003; Shapiro et al.,2005). These authors could identify shape differences in the vertebrae among several primate species and explain some of them as responsive to locomotion. Also, and in spite of her contradictory generalization, Ankel (1967) described differences in the angulation of the dens and the superior articular facets of the axis among humans and other primates.
As observed by other authors (Olivier,1964; Gommery,1996,1997; Aiello and Dean,2002), we also find species differences in the concavity of the superior articular facets (Table 1). Humans exhibit flatter (less concave) facets than apes or baboons. But it is unclear from our data whether the concavity of the facets directly relates to locomotion. Gorilla and Pan, both mainly quadrupedal, also possess relatively flat facets, while Ateles, a frequently brachiating species, has strongly concave articular facets.
Figure 5 displayed the overall atlas shape differences that owe to evolutionary (cross-taxon) and static (within-taxon) allometry. In either, the atlas, especially its arcus posterior, becomes more robust with size and the processus transversus more cranially inclined. Nonhuman static allometry also involves an elongation of the lateral processus. Additionally, we performed these shape regressions on femur head diameter and body size estimates instead of atlas centroid size. The three computations yielded very similar results so that the shape deformations shown in Figure 5 are not only related to atlas size but also to body size. As the cervical vertebrae bear the weight of the head, these allometric shape changes can be construed straightforwardly as adaptations to, or effects of, a heavier head and the resulting more massive nuchal muscles. The muscles connecting the atlas with the head arise at the arcus posterior and the processus transversus of the atlas, so that these structures are mostly affected by allometric increase in robusticity. Whether body size influences locomotion behavior itself, and thus might additionally influence atlas morphology, is still debated (Fleagle and Mittermeier,1980; Doran,1993; Gebo and Chapman,1995; McGraw,1998).
As in nonhuman primates, the processus transversus in the human atlas becomes somewhat enlarged and more cranially oriented with increasing size. But while evolutionary allometry and nonhuman static allometry both indicate an allometric increase of robusticity, human atlas vertebrae seem to become less robust with increasing overall size. This might itself be a result of locomotion: Homo sapiens, as the sole completely upright walking primate, balances the head on the vertebral column instead of bearing it in front of the cervical vertebrae. As Schultz (1942: p. 490) wrote: "very much less force is required to balance the head in adult man than in adults of other primates. The heads of the primates when posed in the ear-eye horizon, are always heavier in front of the occipital condyles than behind them." A quadrupedal primate would thus need to strengthen its nuchal muscles and related bony structures much more than humans would in order to bear a heavier head. The human atlas can thus increase in size without significantly thickening its bony structures. As a consequence, it would appear relatively less robust.
Human Atlas Morphology
As Figures 2 and 4 show, humans exhibit a distinctive atlas morphology. It is not clear whether this distinction is a consequence of our exceptional locomotion pattern or is due to evolutionary processes unrelated to such functional effects. Ankel (1967,2000) argued that the vertebral column of primates is so morphologically and functionally generalized as to permit erectness without major morphological changes in the basic anatomical plan. The erect human column, she argues, is more a functional than a morphological specialization.
For nonhuman primates, we have presented shape differences that are associated with erect vs. quadruped locomotion (Fig. 7). Figure 6 shows the scores along the two shape components that exhibit the highest covariation with locomotion. If the human atlas morphology was an adaptation to its locomotion strategy, then when human specimens are projected onto the figures we would expect the human shape scores to be an extrapolation of the quadrupede vs. erect trend. Contrary to that expectation, we find Homo scores to be intermediate along both components of Figure 6. This finding suggests that perhaps the specific human atlas morphology is not related to locomotion and that the primate atlas already embraces the possibility of bipedalism. Alternatively, human atlas morphology might simply not be predictable from a nonhuman primate sample. It could instead be a unique adaptation not comparable to those for brachiation or vertical climbing.
This discrepancy can also be seen in Figures 4 and 8. While in more erect nonhuman primates the atlas is less robust than in quadruped species, humans exhibit a relatively robust atlas shape, especially at the arcus posterior. Also, the shape and orientation of the facies articularis superior in humans is not as expected from Figure 8. However, both in humans and in the typical erect atlas shape, the processus transversus is oriented more ventrally. Yet we recommend caution in pursuing any adaptationist explanation for this similarity until more functional or experimental data are available (Gould and Lewontin,1979). Also, it is not clear to what degree the differences in atlas shape are inherited and reflect evolutionary adaptations or are instead epigenetic results of different locomotion patterns. Preuschoft et al. (1988), for example, report that Japanese macaques trained to walk bipedally acquire a pronounced lordosis of the lumbar spine due to a relative increase of the ventral lengths of the vertebral bodies.
In summary, there are clear differences in atlas morphology among the nine primate taxa, along with allometric effects that differ between nonhuman primates and humans. While some of these shape differences relate to locomotion, human atlas morphology still cannot be explained by the nonhuman model. Human atlas shape could thus be an adaptation different from that in the brachiating or vertically climbing orthograde primates, or it might be an adaptation unrelated to locomotion behavior.
- Top of page
- Abstract
- Acknowledgements
The authors thank Philipp Gunz, Peter Schmid, Horst Seidler, and T. Bence Viola for helpful suggestions and support in the course of this study, and the museum curators for allowing access to the specimens in their care. They also thank Dennis E. Slice and T. Bence Viola for valuable comments on earlier versions of this manuscript. This study was supported by grant GZ 200.093/I-VI/I/2004 of the Austrian Council for Science and Technology to Horst Seidler.
- Top of page
- Abstract
- Acknowledgements
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text:
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text:
The Law (La legge) (1960)
Critic Consensus: No consensus yet.
Tickets & Showtimes
The Law (La legge) Trailers & Photos
Movie Info
|Art House & International, Drama
|Directed By:
|Jules Dassin
|Written By:
|Jules Dassin, Fran�oise Giroud
|In Theaters:
|May 4, 1960 Wide
|On DVD:
|Sep 28, 2010
Watch it now
as Marietta
as Enrico
as Don Cesare
as Matteo Brigante
as Francesco Brigante
as Donna Lucrezia
as Tonio
as Pizzaccio
as Attilio
as Giulia
as Giuseppina
as Elvira
as Balbo
as Priest
News & Interviews for The Law (La legge)
Critic Reviews for The Law (La legge)
All Critics (5) | Fresh (3) | Rotten (1) | DVD (2)
full of campy star turns, melodramatic confrontations, and relentless moralism
More than its intellectual topics of interest, Dassin's intricately scripted tale is a showcase for his deft gift with the camera.
Audience Reviews for The Law (La legge)
Who's in charge here? Love and lust in an Italian town. Gina Lollobrigida and Melina Mercouri are the main women; Yves Montand and Marcello Mastroianni are the main men. A nice variety of human emotions are shown. Interesting view of an older Italy. I'll be watching this one again.
Discussion Forum
Discuss The Law (La legge) on our Movie forum!


text:
Ted Cruz Responds To Caitlyn Jenner Jab Over Transgender Bathroom Use
Only 8 Major Studio Films In 2015 Had Significant Gay Characters
Hundreds Call On Mississippi Lawmakers To Repeal Anti-Gay Law
'Grace And Frankie,' Starring Jane Fonda, Lily Tomlin, Returns Friday
David Cicilline: Equality Act Needed To Stop States From Discriminating Against LGBT
House Measure Would Undermine Obama's LGBT Executive Order
A group of conservatives led by
Delegate Neil C. Parrott, a Republican opposed to gay rights, have
launched an effort to repeal Maryland's recently approved transgender
protections bill.
Democratic Governor Martin O'Malley has
pledged to sign the Fairness for All Marylanders Act of 2014 (SB
212), which seeks to prohibit discrimination in the areas of
employment, housing, credit and public accommodations on the basis of
gender identity or expression.
The group began collecting signatures
for a ballot initiative at MDPetitions.com, The
Washington Post reported.
At a press conference Tuesday to
announce the effort, Parrott said the group he chairs worries that
the legislation will allow men claiming to be women access to women's
restrooms. The group has dubbed the legislation the "Bathroom
Bill" and its repeal campaign includes imagery of a stick-figure
man climbing over a bathroom stall to reach a stick-figure woman.
"It opens it up to predators, not
necessarily transsexuals, but predators who will take advantage and
go into the opposite-sex bathroom," Parrott told reporters.
"The unintended consequences of this
law demand that we take it to a petition," said Delegate Kathy
Szeliga, also a Republican.
The petition claims that the
legislation "will require businesses to open up their public
facilities … so that men can use the ladies' room and women can use
the men's room." However, an amendment to the bill added to
counter such claims defines gender identity as being "sincerely
held as part of the person's core identity."
On Top Magazine


text:
Sam Shields, B.J. Raji ready to hit open market
Green Bay - Defensive tackle B.J. Raji and cornerback Sam Shields are set to hit the open market on March 8 and nothing seems to be in the way of them moving on to different pastures.
It's not a complete surprise, but the Packers did not use a franchise or transition tag on either free agent, meaning their best hope for retaining them is to up their offer before free agency starts or have the market prove to them that they can't do any better financially outside of Green Bay.
League sources said that deals for Raji and Shields were unlikely before both gain the right to negotiate with other teams Saturday and only a last-second assessment by the Packers would change that. Officially, free agents can't be signed until March 11, so the Packers have five days of exclusive bargaining rights and three more days of pleading their case.
Players can agree to terms with other teams starting on March 8, but they can't sign a contract until the 11th.
This marks the fourth straight year and the seventh time in nine years under general manager Ted Thompson that the Packers have not used the tag.
It would have cost the Packers $9.654 million to franchise Raji and $11.834 million to franchise Shields. Those numbers account for roughly a third of the $35 million in salary cap room the Packers have going into the new football year.
Under franchise rules, a team would have had to give up two first-round picks to the Packers if it wanted to sign one their franchised free agent. The Packers would have had the right to match the offer also. Under the transition tag, which comes at a price tag of about $1 million less than the franchise for each position, the Packers would have had only the right to match an offer.
Raji is arguably one of the top two defensive tackles in this year's free agent class and could conceivably pull in a deal worthy $9 million a year. But after a subpar season, he might find the market drier than he expected and wind up negotiating again with the Packers once his worth is more clearly defined.
Other defensive tackles competing for free agent money are: the New York Giants' Linval Joseph, Dallas' Jason Hatcher, Chicago's Henry Melton, Miami's Randy Starks and Paul Soliai and Minnesota's Kevin Williams.
Don't forget, receiver James Jones tested the free agent market and after he received a cool reception returned to the Packers at a reasonable price on a three-year deal. He is a free agent again this year and will get another crack at the market.
Shields' worth became more defined Monday when Miami CB Brent Grimes agreed to a four-year deal that ESPN.com reported was worth $28 million, including $14 million guaranteed. That deal consists of the same structure as the deal Chicago's Tim Jennings signed in January (half of it guaranteed) and helps define the market for both sides.
Shields is likely to get something more in the Grimes range than the Jennings deal (four years, $22.4 million) because he's only 26. But both Grimes and Jennings had better seasons than Shields and while both are 30, they played as though they have several good years left.
It's possible the open market will be better to Shields than expected since it only takes one team to blow it all up with an unnecessarily inflated deal. Teams like Cleveland, Oakland and Jacksonville have money to burn and they might pay a premium to get Shields to come to their team.
Other corners who will be competing for free agent money are: Tennessee's Alterraun Verner, New England's Aqib Talib, Indianapolis' Vontae Davis and Denver's Dominique Rodgers-Cromartie.
The Packers will be counting on Shields getting no more than Grimes, which would put them back in the running for his services. They still have five days of exclusive negotiating rights with Shields and could raise their offer after seeing the Grimes deal.
Here are the mandatory one-year offers the Packers would have had to make if they tagged either Raji or Shields:
Defensive tackle: $9.654 million
Cornerback: $11.834 million
Defensive tackle: $8.060 million
Cornerback: $10.081 million
Here is a year-by-year look at when the Packers have used a tag since they first became available in 1993:
2014 – No.
2013 – No.
2012 – No.
2011 – No
2010 – DT Ryan Pickett - franchise
2009 – No
2008 – DT Corey Williams - franchise
2007 – No
2006 – No
2005 – TE Bubba Franks, transition
2004 – OT Chad Clifton, franchise
2003 – DT Cletidus Hunt – transition
2002 – No
2001 – No
2000 – No
1999 – WR Antonio Freeman, franchise
1998 – RB Dorsey Levens, franchise
1997 – No
1996 – No
995 – No
1995 – S LeRoy Butler, transition
1994 – OT Ken Ruettgers, TE Jackie Harris, transition
- Sizing up the Packers' draft on the Web
- Crosby at Brookfield Square Mall Sunday for breast cancer event
- Packers land undrafted free agent QB Joe Callahan of Wesley College
- Coach Mike McCarthy on NFL draft
- GM Ted Thompson on the 2016 draft
- Video: Eliot Wolf on Packers draft so far
- Gutekunst: His athleticism stands out more than anything
- Indiana T Jason Spriggs eager to play for Packers
- Twitter reactions to the Packers picking Jason Spriggs at No. 48, Kyler Fackrell at No. 88
- Bills move up to draft ILB Reggie Ragland


text:
Bridging Drug Discovery's Valley of Death...
From Our May 1 Issue: Immunoassays...
Financial Structures Can Determine Where Startups...
From Our May 1 Issue: Where...
An international team of researchers claim stem cell lines with a greater breadth of ethnic genetic diversity must be developed to optimize the use of such cells as research tools and in future therapies. Their analysis of the ethnic background of a number of stem cell lines will form the basis of a new database of human stem cell genetics.
The research will be published in the January issue of Nature Methods in a paper titled "Restricted ethnic diversity in human embryonic stem cell lines."
Scripps Research Institute professor Jeanne Loring, Ph.D., and colleagues used SNP genotyping to evaluate the ethnic origin of 47 human embryonic stem cell lines and five induced pluripotent stem cell lines from laboratories in countries including Korea, Australia, and Finland. They drew on data from the HuMap Project to identify SNPs related to specific ethnic origins.
The results showed that Caucasians were especially well represented among the stem cell lines. East Asian and mixed heritage cell lines were also common. However, there was a notable lack of cell lines representing African ethnicity.
To address this, the researchers used skin cells from an individual of West African Yoruba heritage to create a new stem cell line, which they claim is the first to carry the genetic profile of this ethnic group and thus includes distinct genetic markers for disease risk and drug metabolism.
Dr. Loring and his team suggest greater ethnic diversity in cell lines would allow for more broadly relevant stem cell research and allow the development of safer as well as more effective stem cell therapies and tissue transplants. "Ethnic origin is a critical piece of information that should come with every cell line," Dr. Loring stresses. "Everyone who works with stem cells should be doing this kind of analysis.
"There's not a lot of value in making a new pluripotent stem cell line now unless it has something new to offer. I think that increasing ethnicity and genetic diversity is an important reason for generating new lines."
Do you think that the recent backpedaling on "heart healthy" practices, and other conflicting studies that claim coffee either causes or prevents cancer, confuse the public more than actually help them make informed choices about their diets?
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text:
Premiere: Watch Ital's Video For "Culture Clubs"
Ahron Foster
In this week's Voice cover story, Michaelangelo Matos profiles Daniel Martin-McCormick, the Brooklyn musician currently making waves under the name Ital. In the video for "Culture Clubs," shot live in Cleveland last November by Aurora Halal and premiering on Sound of the City, Martin-McCormick's appeal as both a beatmaker and a live performer is made plain; he churns along with the music, making the energy ripple outward in such a way that makes movement instinctual. Clip below.
ITAL - 'Culture Clubs' live in Cleveland Nov 2011 from Aurora Halal on Vimeo.
In case you want to catch this in action, Ital plays the Knitting Factory on Wednesday, March 21, and the Unsound Festival edition of the Bunker, at Warsaw, on Friday, April 20.
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text:
Lukas Podolski expressed his delight at Arsenal's fourth-place Premier League finish that sealed UEFA Champions League football next season.The striker was Arsene Wenger's marquee signing in July 2012 and he has repaid the faith shown in him by the Frenchman with a double-figure goal tally this season.
120, or even penalties we will see who the winner will be."
Podolski's team-mate and fellow Germany international Per Mertesacker is also unsure who will come out victorious at Wembley when the top two teams in the Bundesliga meet.
"(What is important is) that Bayern and Dortmund show their best possible performances," he said. "Just the way they already have all season long. Lots of pressure on the opposition, quick plays.
"I think we have all been surprised by what is possible for them, especially in the Champions League. I think both teams deserve the win. But there can only be one winner.
"I think there is going to be a great atmosphere surrounding the stadium and all over London."
|S. Harper
|F. Coloccini
|M. Debuchy
|S. Taylor
|M. Yanga-Mbiwa
|H. Ben Arfa
|Y. Cabaye
|J. Gutiérrez
|C. Tioté
|Y. Gouffran
|P. Cissé
|Coach: A. Pardew
|W. Szczęsny
|P. Mertesacker
|B. Sagna
|K. Gibbs
|L. Koscielny
|T. Rosický
|Santi Cazorla
|A. Ramsey
|T. Walcott
|L. Podolski
|Coach: A. Wenger
for Y. Cabaye 57'
for Y. Gouffran 65'
for M. Yanga-Mbiwa 80'
for Arteta 28'
for L. Podolski 78'
for Santi Cazorla 89'
|Ribery tells Guardiola to play him against Atletico
|Vardy wins FWA Player of the Year award
|Hiddink: Title pressure could get to Tottenham
|Messi never cheats, insists Xavi
|Dortmund tell Bayern Hummels fee is non-negotiable
|Welbeck a different animal again - Wenger
|Xavi: Real Madrid played like a small team in Clasico
|Reina backs Atletico to win the double
|Bayern can win Champions League - Santa Cruz
|Perth Glory lure Herd to A-League
|England can´t afford Australia loss - Kewell
|Stankovic: Simeone will coach Inter, but not now
|I had to rotate squad - Pellegrini
|Candreva backs Inzaghi for Lazio job
|I can make fans forget my bad season – Hazard
|MLS Review: Cole sees red in Galaxy stalemate, Valeri stunner lifts Timbers
|Pochettino: Deal announcement can convince Tottenham stars to stay
|Hiddink: Chelsea can´t replace Terry
|Pulisic, Morris picked by Klinsmann in initial Copa squad
|Wenger anticipated worse from protests
|Kante set to have stand named after him
|Furious Mancini slams Inter´s ´worst ever´ first half against Lazio
|Serie A Review: Champions League places locked, Milan play out six-goal thriller
|La Liga Review: Villarreal secure Champions League return
|Lazio 2 Inter 0: Champions League race comes to a Klose
|Koeman delivers Pelle warning
|Morata wants to stay at Juventus, claims Marotta
|Pellegrini disappointed with City defending in defeat
|Iheanacho: City will bounce back from Saints defeat
|´On fire´ Mane salutes Southampton show
|Angers 0 Marseille 1: Batshuayi strike secures Ligue 1 status
|Southampton 4 Manchester City 2: Mane hat-trick blows top-four race open
|Van Gaal calls for video referees after Drinkwater decision
|Schmeichel: I don´t care how we win it
|Montella open to Italy role
|Brocchi pleased with Milan fightback
|Van Gaal defends ´human´ Fellaini over Huth elbow clash
|Ronaldo wants Zidane to stay at Madrid
|Ranieri to miss decisive Tottenham clash
|Rooney disappointed after Leicester draw dents top-four hopes
|Bayern Munich v Atletico Madrid: Hosts seek to vent Bundesliga frustration
|Leicester captain Morgan hails battling Old Trafford point
|AC Milan 3-3 Frosinone: Menez penalty denies strugglers
|Klopp not concerned by Sturridge behaviour
|Manchester United 1 Leicester City 1: Captain Morgan on target as Foxes edge closer to glory
|Allofs: We do not have the same weapons as Bayern
|Guidolin coy on Swansea future
|Allegri happy with win after title celebrations
|Umtiti flattered by Tottenham interest
|Klopp frustrated by lacklustre Liverpool
|Hannover mourn Feierabend
|Godin, Carrasco fit for Bayern clash
|Swansea City 3 Liverpool 1: Ayew double punishes wasteful visitors
|Dybala claims Messi, Ronaldo would score less in Italy
|Juventus 2 Carpi 0: Hernanes, Zaza on target as champions march on
|Ronaldo back in training for Madrid
|Barcelona confirm Bravo calf injury
|Aulas brands Ligue 1 ´championship of Qatar´
|Leicester City at Manchester United: Aiming for Hollywood ending to title push
|I can start for England at Euro 2016 - Sturridge
|Dybala never doubted Juventus
|Aguero celebrates brother´s Independiente debut
|Sturridge rounds on critics, plays down Klopp rift
|Adelaide United 3 Western Sydney Wanderers 1: Reds seal maiden A-League title
|Eriksson: Simeone is an example to follow
|´Where am I?´ - Lovren reveals travel struggles
|´If I have to leave Bayern, then I will´ - Benatia
|Walker ´over the moon´ after Pochettino agrees new deal
|MLS Review: Villa leads City, Impact held
|We´d probably be relegated without Defoe – Allardyce
|Koeman: Leicester will do well in Champions League
|Ronaldo needs rest or could miss Euros - doctor
|United shaped me as a player - Schlupp
|Sanchez, Vidal lead Chile´s Copa squad
|Arsenal´s patience paid off - Cech
|I will know Madrid´s plans - Pellegrini
|Granada 3 Las Palmas 2: Superb comeback boosts hosts´ survival hopes
|Martinez pleased with impact of new faces
|Ligue 1 Review: Lyon, Monaco continue fight for second
|Serie A Review: Fiorentina frustrated, Torino hit five
|Barca keeper Bravo set for tests on calf strain
|Barca will not rely on rivals slipping up - Luis Enrique
|Simeone: I am not a fortune teller
|Neil defiant in defeat
|Real Betis 0 Barcelona 2: Rakitic and Suarez edge champions back ahead in title race
|´Protests more like disappointed love´ - Wenger sorry he cannot keep Arsenal fans happy
|Bundesliga Review: Monchengladbach and Dortmund delay Bayern´s title party
|Arsenal should be doing better, concedes match-winner Welbeck
|Black wants Villa display to build bridges
|Tuchel: Hummels abuse went too far
|Flores laments mixed Hornets performance
|Scottish Premiership Review: Celtic made to wait by Aberdeen
|Ribery a doubt for Bayern´s crunch Atletico clash
|Arsenal 1 Norwich City 0: Substitute Welbeck boosts under-fire Wenger
|Allardyce rues Sunderland nerves after Stoke draw
|Atletico Madrid 1 Rayo Vallecano 0: Griezmann maintains title momentum
|Benitez salutes Newcastle ´heroes´ Townsend and Darlow
|Hughes disappointed by Sunderland penalty call
|Football League Review: Wigan confirm promotion to Championship
|Bilic confident of Europe after West Brom win
|Howe defends players´ efforts after form slump
|Bale can match Ronaldo´s run - Zidane
|Watford poor in Villa win, concedes Deeney
|Hummels slams ´nonsense´ claims over Bayern request
|Zidane hopeful on Ronaldo, Benzema as Madrid give backing to medics
|Pardew rues Cabaye penalty miss in defeat to Newcastle
|West Brom did not get the breaks - Pulis
|Fan protests of no concern to Martinez
|Ramos urges Real Madrid to ´fight to the end´
|Defoe: You haven´t got a choice, I had to score
|Watford 3 Aston Villa 2: Late Deeney double sinks Villa
|Stoke City 1 Sunderland 1: Defoe snatches point but Allardyce´s men slip into bottom three
|Real Sociedad 0 Real Madrid 1: Late Bale winner keeps title challenge alive
|West Brom 0 West Ham 3: Noble brace boosts European hopes
|Newcastle United 1 Crystal Palace 0: Townsend, Darlow heroics boost Benitez
|Everton 2 AFC Bournemouth 1: Baines strike eases pressure on Martinez
|Leno to stay at Leverkusen after buyout clause expires
|Bayern Munich 1 Borussia Monchengladbach 1: Hahn puts title celebrations on hold
|Borussia Dortmund 5 Wolfsburg 1: Fans round on Hummels but title hopes remain
|Watzke won´t give up on keeping Hummels
|Pepe tips Zidane to become top coach
|Allegri and Mancini laud Ranieri´s Leicester heroics
|Allegri still hopes to keep Morata at Juventus
|Mancini praises ´top-quality´ Banega
|´Even Guardiola needed time´ - Brocchi has no regrets over taking AC Milan role
|Willian desperate for next season to arrive
|Mancini: Inter season not a disaster
|Chelsea v Tottenham: Pochettino hopes derby motivation keeps title chase alive
|Crotone not Italy´s Leicester, says director Ursino
|Klopp: Flanagan still faces fitness battle
|Manchester United v Leicester City: Ranieri aiming for Hollywood ending to title push
|Hummels to Bayern is the logical deal - Hitzfeld
|Shaw pushing for United comeback this season
|Sportiello ´proud´ to be linked with Napoli
|Arsenal must change to end title drought - Cazorla
|Tete can´t confirm Bayern interest
|Montero, Valencia set for Copa America Centenario
|Vardy, Mahrez more important than Leicester´s defence - Martinez
|Pellegrini blasts ´unbelievable´ Premier League fixtures
|Standard Liege terminate Valdes´ loan deal
|Hiddink: Leicester will not disappear
|Mata: Leicester can wait a week
|Pizarro, Farfan not included in Peru´s 40-man Copa squad
|Wenger: My loyalty helped build Emirates
|Pochettino eyes Ferguson legacy at Tottenham
|Thompson to leave Melbourne Victory
|New York Red Bulls 4 FC Dallas 0: Hosts make it two in a row
|Blanc hails ´motivated and disciplined´ PSG
|Van Gaal: Easier for Leicester to sign players than United
|No shocks as Paraguay announce provisional Copa squad
|Suarez headlines Uruguay´s Copa America squad
|Sturridge reveals he is ´not Liverpool for life´ and hopes to match Suarez success
|David Luiz, Silva and Marcelo absent from Brazil´s preliminary Copa America squad
|Augsburg 0 Cologne 0: Weinzierl´s men edge closer to survival


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Priest Losing Parish For Being Drunk At A Party?
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Priest Losing Parish For Being Drunk At A Party?
Study Tracks Football Concussions, Resuming Play
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Priest Losing Parish For Being Drunk At A Party?
Holding signs proudly, dozens at St. Charles Borromeo Parish showed how unhappy they were with the Sacramento Diocese`s decision to let go of their Priest Fr. Ron Marcelo. In a letter to parishioners Fr. Ron said he attended a going away party last October where he and staff members drank some wine. Days later, Fr. Ron says he got a letter from Bishop Soto saying he (Fr. Ron) was under the influence of alcohol and was being put on administrative leave. Fr. Ron says he was then asked to attend an evaluation and rehab.
1:20Now watching Up next
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The first concussion study of its kind found youth football players are more likely to return to play less than a day after injury than those in high school and college. (May 1)
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Ida Keeling, who has been running since she was 67, runs in hallways to stay active and train for her record-breaking dash.
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Florida Mom Missing Since Wednesday Sends Investigators Searching
ABC News' David Muir reports the stories people have been buzzing about all week.
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Florida Man Accused of Being a Serial Harasser
A recording of a man allegedly sexually harassing a woman at Target went viral, prompting several other woman to come forward against him.
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Starbucks Faces Legal Trouble for Allegedly Underfilling Drinks
The coffee-chain is accused of intentionally leaving too much room in its cups in order to maximize its profits.
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Tom Angel allegedly composed emails that made insulting jokes toward minorities, an act the chief finds "deeply troubling."
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The parishioners of the Serbian Orthodox Cathedral vow to rebuild the historic landmark.
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SmartScore 99
SmartScore 99
SmartScore 86
SmartScore 92
Here's the question you clicked on:
Which of the following mechanisms moved slower, the worm and wheel or the rack and pinion?
Worm and wheel is slower because for each full turn of the worm shaft only one tooth of the wheel is displaced. In the case of the rack and pinion, the full turn of the pinion displaces an equivalent amount of teeth in the rack.


text: Heil Bush as he leads us down the hershey highway. Gas to go to $3 and stocks to crash. But hey if we only agree to put our social security into the market we will all be saved. He feeds his Saudi friends with our blood and guts. Heil Hitler!!!! You Americans are so pathetic and stupid.


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Affiliation will give area residents access to UPMC's world-class health care in their own backyard.Read More>>
Pitt's GSPH launches Cura Zika, an international alliance to perform much-needed research addressing the Zika epidemic.Read More>>
Pitt has chosen a renowned faculty member as the next chair of the Department of Physical Medicine and Rehabilitation.Read More>>
Pitt researchers have discovered more than 500 new protein-protein interactions associated with genes linked to schizophrenia.Read More>>
Thursday, May 5, 2016
UPMC, Lycoming County health system join forces
Casey visits Pittsburgh hospital, aims to reduce number of babies born into opioid dependence
Friday, May 6, 2016
Pitt Researchers Partner With Brazilian Government On Zika Research Thursday, May 5, 2016FDA brings e-cigarettes under federal authority Thursday, May 5, 2016
Earnhardt willing for his brain to be a matter of science
Richard Zimmerman, M.D., M.P.H., professor of Pitt's Department of Family Medicine and Graduate School of Public Health, has been awarded the Curtis G. Hames Research Award by the Society of Teachers of Family Medicine. Read more
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Sheviock Holiday Cottages
Please find below a selection of self catering holiday cottages to rent in or near Sheviock.Read More
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We open tomorrow at 9am. You can also send us an e-mail or request a callback.Request a callback
Please find below a selection of self catering holiday cottages to rent in or near Sheviock.Read More
Your search returned 49 cottages.
Single-storey, stone barn conversion in rural Cornwall near Liskeard. Open plan living area, modern styling and traditional character features. WiFi. Liskeard 3 miles. (Ref. 5240)
7 nights from £260Book Now
Traditional, Cornish stone-built barn conversion. Warm and cosy family accommodation in the countryside, ten miles from coast. WiFi. Charming rural views. Liskeard 3 miles. (Ref. 5260)
7 nights from £347Book Now
Lovely, stone-built cottage in peaceful Cornish village of St Keyne. Perfect romantic hideaway. Single track rail line to Looe nearby. Liskeard 3 miles. (Ref. 2470)
7 nights from £248Book Now
A charming, stone-built cottage in the peaceful Cornish village of St Keyne, 3 miles from Liskeard. Cosy, comfortable accommodation. Easy access to South Coast. Looe 5 miles. (Ref. 1678)
7 nights from £265Book Now
A pretty character cottage on the banks of the River Tamar. Sun deck overlooking the river. Use of moorings. Tavistock 6 miles. (Ref. 16511)
7 nights from £302Book Now
Victorian barn conversion set in an idyllic, rural location on a non-working farm, midway between Bodmin and Dartmoor. Ideal for walkers. Open plan living room with beams, a quiet spot for wildlife spotting. WiFi. Callington 2 miles. (Ref. 918087)
7 nights from £292Book Now
Delightful south-facing bungalow located in a quiet road in the village of St Keyne. Two bedrooms. WiFi. Off road parking on driveway. Enclosed garden. Liskeard 3 miles. (Ref. 7406)
7 nights from £265Book Now
A wonderful detached cottage situated in the Cornish village of Harrowbarrow. Set in extensive gardens with outdoor solar heated swimming pool and wildlife garden. Tavistock 5 miles. (Ref. 20763)
7 nights from £376Book Now
Semi-detached woodland lodge with countryside views, on Honicombe Manor holiday park. Spacious accommodation. Free Country Club leisure pass. Tavistock 5 miles. (Ref. 5147)
7 nights from £335Book Now
A delightful Alpine lodge in a quiet park in Tamar Valley AONB. Open plan living area with balcony and far-reaching views. Seasonal on-site indoor and outdoor heated swimming pools, plus tennis courts. Gunnislake 2.5 miles. (Ref. 17180)
10% off - Sat 14th May to 21st May 2016
7 nights from £361Book Now


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Doctor-patient ties worsening across India?Mumbai: Association of medical consultants (AMC) trustee Dr Niranjan Agarwal's reaction to a Sion hospital's resident doctor being roughed up on Wednesday says a lot about the doctor-patient relationship. "Every death or complication is doctor's ... Times of India, 1 month ago
Experts call for MCIs radical reforming
Irish Sun (ANI) Friday 1st April, 2016 Washington D.C, Apr 1 (ANI): A team of leading doctors has welcomed a landmark report calling for radical reform of the Medical Council of India (MCI). Dr Sanjay Nagral, a consultant surgeon at Jaslok ...IrishSun.com, 1 month ago Top docs back House panel proposal to overhaul MCI Times of India, 1 month ago Medical experts call for MCI's radical restructuring Asian Age, 1 month ago British Medical Journal calls for radical revamp of MCI The Hindu, 1 month ago
Leopard mysteriously drowns in well with parapet wall
Chandrapur: Decomposed carcass of a sub-adult leopard was found in a farm well in suspicious conditions, on the outskirts of Kothari village on Wednesday morning. While veterinary doctors concluded the beast died of drowning, the mystery is how the ...Times of India, 1 month ago
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The Graduates 2011: Doug Stewart
Originally from Glastonbury where he did his foundation, Doug Stewart studied graphic design at Kingston University and left with a healthy appreciation for collaboration and broad skill set that far outweighs his graduate status. A fascinating exploratory type project, Regausser, uses the disruptive qualities of magnets, controlled and applied creatively to television sets to create new forms and emphasising Doug's love of all things multi-disciplinary and multi-media.
This "pick & mix approach that can be used to create something fresh and new" is just one way he challenges himself. The aforementioned collaborative facet that Kingston thrives on, led him into video works with fellow graduate, Scott Taylor. While a personal piece, Night Tracks, integrates music, design, and illustration into a public environment. Inspired by nocturnal suburbia, "tracks are temporarily illustrated within the night-time environments that influenced them."
If your portfolio was on fire, and you could only save one piece/project, which would you choose, and why?
Wow, well this year I've worked on so many different projects of which I'm proud that it's hard to choose just one. But A3 re-prints come pretty cheap, so I'd probably try and save the box – it took a long while to apply my vinyl name sticker to it.
If you could collaborate with another artist/designer (or a number of artists/designers) to make a piece of work, who would you work with and what would you make?
I'm lucky to be in a situation where I have come from a course that is heavily structured around collaboration, and from that, have developed strong working relationships with many fellow course members. It would be exciting to be able to collaborate with some of my Kingston course mates again in the future. But if they're not available, then working with Heston Blumenthal would be cool. I've come up with a concept for a new food product that we might be able to collaborate on. It's half way between a Pasty and a Quiche and I'm branding it a 'Pastiche.' If you're reading this Heston, I'm happy to pay your travel costs.
What was your finest moment at art school?
Our team winning the pub quiz on our year 2 Graphics & Illustration field trip to San Francisco ranks up there pretty highly. Besides that, 'Step Trip Step,' our recent degree show that we put on ourselves in London was a massively proud moment. And also having my degree show wall piece featured in D&AD's New Blood in July. That's a top 3; I cheated.
We believe it was the Jonas brothers who once said "we're the kids of the future." How, if at all, do you relate to that?
I rarely agree with the Jonas Brothers' wildly outspoken opinions, but I think in this case, I can relate to what they're trying to say. It's exciting to think that alongside other graduates, we are now in a position where we can start to contribute to the kind of work that will define, shape and change the creative industry from this point onwards.
Can you give us ONE prediction about you and your work for the next year?
Ideally, I'll be working or interning in a situation where I can continue to create work that makes me happy, alongside developing my skills. (As well as making enough money to live off the back of it.) This is not so much a prediction, but more of a pipe dream! I guess the fact that I don't know where I'll be in 12 months time is what makes this post-university stage so exciting right now.
- Submit Saturdays: photographer and filmmaker Harry Israelson's bright, smart portfolio
- May Diary: where to go and what to see this month
- Crisp and vibrant design work from ECAL graduate Clement Rouzaud
- Portuguese illustrator Tiago Galo's plump little characters are oddly charming
- Matthew Butcher launches the Flood House that will travel around the Thames Estuary
- Haunting train-simulator-based animation by Jack Featherstone for Occult Orientated Crime
- Philip Coppola spends nearly 40 years illustrating New York City's Subway Stations
- LA studio Laundry creates amazing warped Simpsons idents for American channel FX
- Design Bridge creates new harp icon for Guinness
- Winning design for Tokyo 2020 Olympics unveiled
- Prince: 1958-2016
- Milton Glaser creates new look for Brooklyn Brewery


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The deadline for this year's Toshiba America Foundation's Science and Math Improvement Grant for K-6 is looming. Educators have until Oct. 1 to apply for the grants, which award up to $1,000 to fund science or math projects. The Toshiba America Foundation (TAF) is urging educators to apply, saying the program "is very easy to access for science and math teachers."
The Science and Math Improvement Grant awards schools for "programs and activities that improve teaching and learning in science and mathematics" in K-12 education. K-6 teachers can receive $1,000 mini-grants for math and science projects. Teachers of grades 7 through 12 can receive grants of up to $5,000. The deadline for K-6 grant applications is Oct. 1. Teachers of grades 7 through 12 can apply at any time during the calendar year. Teachers can also apply for grants larger than $5,000. Such applications will be reviewed twice per year: Feb. 1 and Aug. 1.
The Toshiba America Foundation has so far granted some $9 million to educators since its inception, supporting about 1,600 schools in the United States, Canada, and Mexico. More information about the grant program can be found here.
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Students need a broad set of skills to be successful in today's world. To better help them prepare, K-12 districts throughout the country are taking a blended approach to achieve personalized learning. Making that transition successfully starts with envisioning what learning should look like and how to create authentic learning experiences. Download this white paper to learn how two school districts, Cabarrus County Schools in North Carolina and Lake Travis Independent School District in Texas are currently making that transition.
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Chefn Pk.2 Eco Crock Charcoal Filters 401421001
|Chefn Pk.2 Eco Crock Charcoal Filters 401421001Made for use with Chef'n Meringue & Black Eco Crock Compost BinSet of 2 natural charcoal filters
Chef'n Switchit-All Purpose Spatula-Cherry 103 656 005
|The Chef'n Switchit All Purpose Cherry Spatula is a 11.5" scraper and stirer made from a steel core wrapped in heat and stain resistant silicone. This all purpose tool features a large spatula on one end and a small, narrow spatula on the other. The Silicone is heat-safe to 650° F. and is excellent for scraping, folding and stirring. It will not scratch nonstick or any other cookware and is top rack dishwasher safe. Measures 13.5" l x 3.5" w x 0.5" h.
Chefn Eco Crock Compost Bin-Meringue/Black 401420120
|Chefn Eco Crock Compost Bin-Meringue/Black 401420120Lid features sprout-shaped handle and ventilation holes to promote airflowHas natural charcoal filter in the lid to help absorb odors. Holds 3.5 quarts Has a removable inner plastic liner
Chef'n G'rabbit Jr Salt+pepper Set
|As if the original G'Rabbit salt and pepper grinders weren't already cute enough, Chef'n went ahead and created mini-magnetic versions! Introducing the G'Rabbit Jrs: adorable, convenient and full of flavor, they're the ultimate kitchen accessory. A rare earth magnet on the backside of the G'Rabbit Jrs allows the grinders to attach to any metal surface offering a creative storage solution. Set of two; includes one G'Rabbit Jr Pepper Grinder and one G'Rabbit Salt Grinder; filled with either Mediterranean Sea Salt or a Global Pepper Medley.
Chefn Dual Pepper Ball-Black/Clear 101182001
|Chefn Dual Pepper Ball-Black/Clear 101182001Dual grinder with 2-in-1 design Twist the handles to select salt or pepper Squeeze handles to grind. To refill, twist off the base. Adjustable grinding mechanism Comes prefilled with sea salt and peppercorns.
Chef'n Garlic Machine 102474009
|Garlic press, Multi-clover loader, Efficient mechanism for mincing, Pop on bottom cap for refrigerator storage, Top rack dishwasher safe
Chef'n Lemon-fresh Force Citrus Juicer
|The FreshForce Citrus Juicer is perfect for pressing juice from halved lemons and limes. Use it for making margaritas, lemonade or adding that extra citrus zing to desserts and entrees. Unlike other juicers, the FreshForce Citrus Juicer has an extra gear mechanism that increases juicing power and minimizes hand strain. Easy to squeeze handle. Dishwasher safe. Lemon colored finish. By Chef'n. Chef'n began over twenty years ago based on the principle that kitchen products could be a lot smarter, easier and more fun. Right from day one, Chef'n found an audience hungry for its brand of tasteful ingenuity. Today that legacy lives on with the precision refinements to acclaimed inventions, new technologies, design techniques and entirely new products.
Chefn Arugula-Fresh Force Lime Juicer 102415001
|Chefn Arugula-Fresh Force Lime Juicer 102415001 Dual-gear mechanism squeezes 20% more juice than traditional juicers To juice, insert lime cut side down and squeeze. Top rack dishwasher safe.
Chef'n Veggisteam Cherry Vegetable Steamer
|With its innovative construction and expanded surface area, the The Chef'n 8.5" VeggiSteam Cherry is a steaming revolution. This steamer insert has a silicone body with a nylon core and is flexible to fit snug in any pan size smaller than 9" in diameter. It is 650°F (340°C) heat resistant and will not stain or retain food tastes and odors. All VeggiSteams are top rack dishwasher safe and safe to use in the microwave. Cherry color. Measures 8.5" l x 8.5" w x 1.25" h.
Chef'n Sweet Spot Ice Cream Sandwich Maker 107 821 251
|The Chef'n Sweet Spot Ice Cream Sandwich Maker helps you make delicious ice cream sandwiches at home! The kit includes 2 silicone baking sheets, a shaping and freezing mold and a spreader tool. The kit can make 4 ice cream sandwiches at the same time and is top rack dishwasher safe. Measures 9" l x 6" w x 1.25" h.
Chef'n Stemgem Strawberry Stem Remover
|Remove strawberry stems with ease with the StemGem strawberry stem remover from Chef'n. It's easy to use; simply insert the StemGem into strawberry, twist and remove the strawberry stem. It removes the stem and leaves the strawberry intact! It's great for making strawberry shortcake and fruit salads. And, with the fun strawberry design, it's easy to find in the drawer. Claw made of stainless steel. Top-rack dishwasher safe.
Chef'n Mini Magnetic Pepper Ball Set
|This set of Chef'n Mini Magnetic salt and pepper grinders combines a whimsical bunny design with easy-to-use functionality. To activate them you simply squeeze the "ears" together. They are constructed of sturdy, clear acrylic with the handles having soft rubber grip pads. Use the easy slide refill window to replenish the salt and pepper supplies and the strong magnets to always keep them handy on the front of the fridge or stove.6-1/2" H x 5-1/2" W l
Chef'n Potato Ricer-fresh Force
|The FreshForce Potato (Ricer) Press makes prepping mashed potatoes a breeze! Unlike other presses, the FreshForce Potato (Ricer) Press has an extra gear mechanism that increases ricing power and minimizes hand strain. Easy to squeeze handles. Stainless steel basket. Dishwasher safe. By Chef'n. Chef'n began over twenty years ago based on the principle that kitchen products could be a lot smarter, easier and more fun. Right from day one, Chef'n found an audience hungry for its brand of tasteful ingenuity. Today that legacy lives on with the precision refinements to acclaimed inventions, new technologies, design techniques and entirely new products.
Chef'n Flexicado Avocado Slicer
|The Flexicado, flexible avocado slicer, slices avocados quickly and easily. It's perfect for prepping guacamole! It fits any size avocado; simply squeeze the raised finger-grip to fit Flexicado onto any size avocado. Features an extra fin for more consistent slices. Top-rack dishwasher safe. By Chef'n. Chef'n began over twenty years ago based on the principle that kitchen products could be a lot smarter, easier and more fun. Right from day one, Chef'n found an audience hungry for its brand of tasteful ingenuity. Today that legacy lives on with the precision refinements to acclaimed inventions, new technologies, design techniques and entirely new products.
Chef'n Quickpit Cherry Pitter
|Stop biting down on hard cherry pits! This hand-held cherry pitter gets rid of cherry and olive pits in seconds. Just insert a cherry or olive into the nook, pull the trigger, and watch the pit pop right out! This gadget is perfect for kids since it does not require a blade.
Chef'n Switchit-wooden Slotted Turner-arugula
|Great for stirring and scraping food while it cooksAttractive wood design is safe to use on non-stick cookware650?F (340?C) heat and stain resistant silicone spatula endHand wash recommended
Chefn Pepper Ball-Black/Clear 101235001
|Chefn Pepper Ball-Black/Clear 101235001Like the classic pepper ball, only betterReinforced handlesAdjustable grinding mechansim Steel rasp To refill, twist off top.Comes prefilled with fresh peppercorns.
Chef'n Clear Relish Pepper Grinder 101299002
|Grind pepper quickly and efficiently with this Chef'n Clear Relish Pepper Grinder. It features a twist-top design and ceramic rasp to grind pepper easily. Adjust the coarseness of your grind by turning the adjustment dial on the bottom of the grinder. To refill, remove the top.
Chef'n Clear Relish Salt Grinder 101300002
|Grind salt quickly and efficiently with this Chef'n Clear Relish Salt Grinder. It features a twist-top design and ceramic rasp to grind salt easily without worry of corrosion. Adjust the coarseness of your grind by turning the adjustment dial on the bottom of the grinder. To refill, remove the top.
Chef'n Zeel Peel 102546173
|The Chef'n Zeel Peel makes quick work of peeling oranges. Just use the sharp hook to score the skin, then lift it back with the pointed corner, then slide the edge through the opening and pull down. It's that simple! Top rack dishwasher safe.
Chef'n PinPair Silicone & French Rolling Pin Set 103 551 005
|The Chef'n PinPair is a silicone baker's pin and a French wooden rolling pin in one. With a simple twist you can remove the wooden French-style rolling pin from the silicone pin. Top rack dishwasher safe.
Chef'n Emulstir Salad Dressing Mixer 104555011
|For a healthy alternative to high-calorie salad dressings, Chef?n introduces the new and improved Emulstir. This emulsifier and salad dressing mixer is perfect for whipping up healthy homemade dressing in no time. To use, simply fill with your favorite ingredients. Squeeze the handle to activate the mixing lever and emulsify. Ingredients are smoothly and evenly blended. Pull up integrated cap for easy and clean pouring. Close the cap and place in refrigerator for later.
Chef'n Pastry Pen 103 549 005
|The Chef'n Pastry Pen is an all-in-one cupcake baking and decorating tool.The Pastry Pen includes everything you need to create, fill and decorate your homemade treats.
Chef'n Veggisteam Arugula Vegetable Steamer
|The Chef'n 8.5" VeggiSteam Arugula is a steaming revolution! This steamer insert has a flexible silicone body with a nylon core that can fit snug in any pan size smaller than 9" in diameter. It is 650°F (340°C) heat resistant and safe to use on the stovetop, in the oven or michrowave. The VeggieSteam will not stain or retain food tastes and odors and is top rack dishwasher safe. Arugula color. Measures 8.5" l x 8.5" w x 1.25" h.
Chef'n Switchit-Mini Spoon-Arugula 103 672 011
|The Chef'n Switchit Arugula Mini Spoon is a handy double-sided spoon that is stain resistant and heat safe up to 650° F. The narrow shape can reach into narrow blenders, jars, and containers unlike any other kitchen tool. It's also great for scraping mixer attachments. It is safe for stovetop use and will not scratch non-stick cookware.
Chef'n Switchit-Mini Spoon-Cherry 103 672 005
|The Chef'n Switchit Cherry Mini Spoon is a 2-In-1 narrow spoon tool with two usable work ends that are ideal for a variety of jobs at any temperature. The outer silicone construction is flexible, stain-resistant and heat safe to 650° F. It's great for getting into tight spaces including jars and mixer paddles. Top rack dishwasher safe.
Chef'n A Jar Jar Opener Cherry 103 661 005
|The Chef'n Ajar Jar Opener adjusts to any size lid and locks on to provide leverage for easy opening. The comfortable red handle helps you grip and has an arrow imprinted so you know which way to turn.
Chef'n Garlic Zoom Assorted 102 680 229
|Keep the flavor in your food and the smell off your hands with the Chef'n Garlic Zoom. This redesigned model is a snap to use, just put peeled garlic inside and roll! The stainless interior blades with chop the garlic the way you want it. The new Garlic Zoom features improved gearing for easier rolling, bigger finger holds for safer blade removal and sturdier construction. And when done, your Garlic Zoom is top rack dishwasher safe for easy cleaning. Measures 2.3" lx 2" w x 2.9" h. **Color will vary.
Chef'n Bramble Berry Basket and Colander, 1-Quart, Cherry
|The Chef'n Bramble Berry Basket is perfect for rinsing, storing and serving fresh berries. The inner basket removes and acts as a berry colander for easy washing. Use in the garden, take along to the store or great for farmer's markets. Great for the recycle-conscious consumer. Containers are stackable which make storing easy. The Chef'n Berry Basket is BPA-free and top-rack dishwasher safe. Measures 4.5" l x 4.5" w x 4" h.
Chef'n Pop Top Microwave Popcorn Popper, Cherry
|The Chef'n Poptop Popcorn Popper is constructed from high-heat resistant silicone, and has sides that fold in to become a lid, which then unfold as kernels pop. The Poptop will hold up to 10 cups of popped popcorn. Simple to store and top-rack dishwasher safe.