Thoracic vertebral count and thoracolumbar transition in Australopithecus afarensis.

The evolution of the human pattern of axial segmentation has been the focus of considerable discussion in paleoanthropology. Although several complete lumbar vertebral columns are known for early hominins, to date, no complete cervical or thoracic series has been recovered. Several partial skeletons have revealed that the thoracolumbar transition in early hominins differed from that of most extant apes and humans. Australopithecus africanus, Australopithecus sediba, and Homo erectus all had zygapophyseal facets that shift from thoracic-like to lumbar-like at the penultimate rib-bearing level, rather than the ultimate rib-bearing level, as in most humans and extant African apes. What has not been clear is whether Australopithecus had 12 thoracic vertebrae as in most humans, or 13 as in most African apes, and where the position of the thoracolumbar transitional element was. The discovery, preparation, and synchrotron scanning of the Australopithecus afarensis partial skeleton DIK-1-1, from Dikika, Ethiopia, provides the only known complete hominin cervical and thoracic vertebral column before 60,000 years ago. DIK-1-1 is the only known Australopithecus skeleton to preserve all seven cervical vertebrae and provides evidence for 12 thoracic vertebrae with a transition in facet morphology at the 11th thoracic level. The location of this transition, one segment cranial to the ultimate rib-bearing vertebra, also occurs in all other early hominins and is higher than in most humans or extant apes. At 3.3 million years ago, the DIK-1-1 skeleton is the earliest example of this distinctive and unusual pattern of axial segmentation.

Human torque is not present in chimpanzee brain.

We searched for positional brain surface asymmetries measured as displacements between corresponding vertex pairs in relation to a mid-sagittal plane in Magnetic Resonance (MR) images of the brains of 223 humans and 70 chimpanzees. In humans deviations from symmetry were observed: 1) a Torque pattern comprising right-frontal and left-occipital "petalia" together with downward and rightward "bending" of the occipital extremity, 2) leftward displacement of the anterior temporal lobe and the anterior and central segments of superior temporal sulcus (STS), and 3) posteriorly in the position of left occipito-temporal surface accompanied by a clockwise rotation of the left Sylvian Fissure around the left-right axis. None of these asymmetries was detected in the chimpanzee, nor was associated with a sex difference. However, 4) an area of cortex with its long axis parallel to the olfactory tract in the orbital surface of the frontal lobe was found in humans to be located higher on the left in females and higher on the right in males. In addition whereas the two hemispheres of the chimpanzee brain are equal in extent in each of the three dimensions of space, in the human brain the left hemisphere is longer (p = 3.6e-12), and of less height (p = 1.9e-3), but equal in width compared to the right. Thus the asymmetries in the human brain are potential correlates of the evolution of the faculty of language.

Skill not athleticism predicts individual variation in match performance of soccer players.

Just as evolutionary biologists endeavour to link phenotypes to fitness, sport scientists try to identify traits that determine athlete success. Both disciplines would benefit from collaboration, and to illustrate this, we used an analytical approach common to evolutionary biology to isolate the phenotypes that promote success in soccer, a complex activity of humans played in nearly every modern society. Using path analysis, we quantified the relationships among morphology, balance, skill, athleticism and performance of soccer players. We focused on performance in two complex motor activities: a simple game of soccer tennis (1 on 1), and a standard soccer match (11 on 11). In both contests, players with greater skill and balance were more likely to perform better. However, maximal athletic ability was not associated with success in a game. A social network analysis revealed that skill also predicted movement. The relationships between phenotypes and success during individual and team sports have potential implications for how selection acts on these phenotypes, in humans and other species, and thus should ultimately interest evolutionary biologists. Hence, we propose a field of evolutionary sports science that lies at the nexus of evolutionary biology and sports science. This would allow biologists to take advantage of the staggering quantity of data on performance in sporting events to answer evolutionary questions that are more difficult to answer for other species. In return, sports scientists could benefit from the theoretical framework developed to study natural selection in non-human species.

Midsagittal Brain Variation among Non-Human Primates: Insights into Evolutionary Expansion of the Human Precuneus.

The precuneus is a major element of the superior parietal lobule, positioned on the medial side of the hemisphere and reaching the dorsal surface of the brain. It is a crucial functional region for visuospatial integration, visual imagery, and body coordination. Previously, we argued that the precuneus expanded in recent human evolution, based on a combination of paleontological, comparative, and intraspecific evidence from fossil and modern human endocasts as well as from human and chimpanzee brains. The longitudinal proportions of this region are a major source of anatomical variation among adult humans and, being much larger in Homo sapiens, is the main characteristic differentiating human midsagittal brain morphology from that of our closest living primate relative, the chimpanzee. In the current shape analysis, we examine precuneus variation in non-human primates through landmark-based models, to evaluate the general pattern of variability in non-human primates, and to test whether precuneus proportions are influenced by allometric effects of brain size. Results show that precuneus proportions do not covary with brain size, and that the main difference between monkeys and apes involves a vertical expansion of the frontal and occipital regions in apes. Such differences might reflect differences in brain proportions or differences in cranial architecture. In this sample, precuneus variation is apparently not influenced by phylogenetic or allometric factors, but does vary consistently within species, at least in chimpanzees and macaques. This result further supports the hypothesis that precuneus expansion in modern humans is not merely a consequence of increasing brain size or of allometric scaling, but rather represents a species-specific morphological change in our lineage.

Qualitative and Quantitative Analysis of Primary Neocortical Areas in Selected Mammals.

The present study focuses on the relationship between neocortical structures and functional aspects in three selected mammalian species. Our aim was to compare cortical layering and neuron density in the projection areas (somatomotor, M1; somatosensory, S1; auditory, A1; and visual, V1; each in a wider sense). Morphological and design-based stereological analysis was performed in the wild boar (Sus scrofa scrofa) as a representative terrestrial hoofed animal (artiodactyl) and the common dolphin (Delphinus delphis) as a highly derived related aquatic mammal (cetartiodactyl). For comparison, we included the human (Homo sapiens) as a well-documented anthropoid primate. In the cortex of many mammals, layer IV (inner granular layer) is the main target of specific thalamocortical inputs while layers III and V are the main origins of neocortical projections. Because the fourth layer is indistinct or mostly lacking in the primary neocortex of the wild boar and dolphins, respectively, we analyzed the adjacent layers III and V in these animals. In the human, all the three layers were investigated separately. The stereological data show comparatively low neuron densities in all areas of the wild boar and high cell counts in the human (as expected), particularly in the primary visual cortex. The common dolphin, in general, holds an intermediate position in terms of neuron density but exhibits higher values than the human in a few layers. With respect to the situation in the wild boar, stereological neuron counts in the dolphin are consistently higher, with a maximum in layer III of the visual cortex. The extended auditory neocortical field in dolphins and the hypertrophic auditory pathway indicate secondary neurobiological adaptations to their aquatic habitat during evolution. The wild boar, however, an omnivorous quadruped terrestrial mammal, shows striking specializations as to the sensorimotor neurobiology of the snout region.

Synaptosomal lactate dehydrogenase isoenzyme composition is shifted toward aerobic forms in primate brain evolution.

With the evolution of a relatively large brain size in haplorhine primates (i.e. tarsiers, monkeys, apes, and humans), there have been associated changes in the molecular machinery that delivers energy to the neocortex. Here we investigated variation in lactate dehydrogenase (LDH) expression and isoenzyme composition of the neocortex and striatum in primates using quantitative Western blotting and isoenzyme analysis of total homogenates and synaptosomal fractions. Analysis of isoform expression revealed that LDH in synaptosomal fractions from both forebrain regions shifted towards a predominance of the heart-type, aerobic isoform LDH-B among haplorhines as compared to strepsirrhines (i.e. lorises and lemurs), while in the total homogenate of the neocortex and striatum there was no significant difference in LDH isoenzyme composition between the primate suborders. The largest increase occurred in synapse-associated LDH-B expression in the neocortex, with an especially remarkable elevation in the ratio of LDH-B/LDH-A in humans. The phylogenetic variation in the ratio of LDH-B/LDH-A was correlated with species-typical brain mass but not the encephalization quotient. A significant LDH-B increase in the subneuronal fraction from haplorhine neocortex and striatum suggests a relatively higher rate of aerobic glycolysis that is linked to synaptosomal mitochondrial metabolism. Our results indicate that there is a differential composition of LDH isoenzymes and metabolism in synaptic terminals that evolved in primates to meet increased energy requirements in association with brain enlargement.

Proximal metatarsal articular surface shape and the evolution of a rigid lateral foot in hominins.

This study quantifies the proximal articular surface shape of metatarsal (MT) 4 and MT 5 using three-dimensional morphometrics. Humans and apes are compared to test whether they have significantly different shapes that are skeletal correlates to comparative lateral foot function. In addition, shod and unshod humans are compared to test for significant differences in surface shape. The MT 4 fossils OH 8, Stw 628, and AL 333-160, and the MT 5 fossils AL 333-13, AL 333-78, OH 8, and Stw 114/115 are compared with humans and apes to assess whether they bear greater similarities to humans, which would imply a relatively stable lateral foot, or to apes, which would imply a flexible foot with a midfoot break. Apes have a convex curved MT 4 surface, and humans have a flat surface. The MT 4 fossils show greater similarity to unshod humans, suggesting a stable lateral foot. Unshod humans have a relatively flatter MT 4 surface compared with shod humans. There is much overlap in MT 5 shape between humans and apes, with more similarity between humans and Gorilla. The fossil MT 5 surfaces are generally flat, most similar to humans and Gorilla. Because of the high degree of shape overlap between humans and apes, one must use caution in interpreting lateral foot function from the proximal MT 5 surface alone.

The enamel-dentine junction in the postcanine dentition of Australopithecus africanus: intra-individual metameric and antimeric variation.

We used micro-computed tomography and virtual tools to study metric and morphological features at the enamel-dentine junction and on the outer enamel surface in the postcanine dentition of an exceptionally well-preserved maxilla and mandible of an early hominin. The fossil, Sts 52 from Sterkfontein, South Africa, is attributed to Australopithecus africanus and is about 2.5 million years old. For comparative purposes in this exploratory study, we also used micro-computed tomography to analyse the dentition of a common chimpanzee (Pan troglodytes), a pygmy chimpanzee (Pan paniscus) and three extant humans. Metameric variation of the 3D enamel-dentine junction in the two chimpanzee mandibles was much smaller than in extant humans. Variation in metameric shape was high and complex. Notably, the mandibular metameric variation in extant humans can be greater within individuals, as compared with variation between individuals, with differences in shape appearing greater for M2 compared with M1. We recommend the use of a new approach in which individual metameric variation is systematically assessed before making inferences about differences between fossil hominin species. The fossil hominin examined in this study showed a metameric pattern of mandibular variation in shape that was comparable to the pattern seen in two chimpanzees. This degree of metameric variation appeared relatively small compared with the much larger patterns of variation observed within and between extant humans.

Pyramidal neurons in the septal and temporal CA1 field of the human and hedgehog tenrec hippocampus.

The present study examines comparatively the cellular density of disector-counted/Nissl-stained CA1 pyramidal neurons and the morphometric characteristics (dendritic number/length, spine number/density and Sholl-counted dendritic branch points/20 microm) of the basal and apical dendritic systems of Golgi-impregnated CA1 neurons, in the septal and temporal hippocampus of the human and hedgehog tenrec brain. The obtained results indicate that in both hippocampal parts the cellular density of the CA1 pyramidal neurons is lower in human than in tenrec. However, while the human pyramidal cell density is higher in the septal hippocampal part than in the temporal one, in the tenrec the density of these cells is higher in the temporal part. The dendritic tree of the CA1 pyramidal cells, more developed in the septal than in temporal hippocampus in both species studied, is in general more complex in the human hippocampus. The basal and the apical dendritic systems exhibit species related morphometric differences, while dendrites of different orders exhibit differences in their number and length, and in their spine density. Finally, in both species, as well as hippocampal parts and dendritic systems, changes of dendritic morphometric features along ascending dendritic orders fluctuate in a similar way, as do the number of dendritic branch points in relation to the distance from the neuron soma.

The evolution of the complex sensory and motor systems of the human brain.

Inferences about how the complex sensory and motor systems of the human brain evolved are based on the results of comparative studies of brain organization across a range of mammalian species, and evidence from the endocasts of fossil skulls of key extinct species. The endocasts of the skulls of early mammals indicate that they had small brains with little neocortex. Evidence from comparative studies of cortical organization from small-brained mammals of the six major branches of mammalian evolution supports the conclusion that the small neocortex of early mammals was divided into roughly 20-25 cortical areas, including primary and secondary sensory fields. In early primates, vision was the dominant sense, and cortical areas associated with vision in temporal and occipital cortex underwent a significant expansion. Comparative studies indicate that early primates had 10 or more visual areas, and somatosensory areas with expanded representations of the forepaw. Posterior parietal cortex was also expanded, with a caudal half dominated by visual inputs, and a rostral half dominated by somatosensory inputs with outputs to an array of seven or more motor and visuomotor areas of the frontal lobe. Somatosensory areas and posterior parietal cortex became further differentiated in early anthropoid primates. As larger brains evolved in early apes and in our hominin ancestors, the number of cortical areas increased to reach an estimated 200 or so in present day humans, and hemispheric specializations emerged. The large human brain grew primarily by increasing neuron number rather than increasing average neuron size.

The lumbar spine in Neanderthals shows natural kyphosis.

Nowadays, lumbar spondylosis is one of the most frequent causes of lower back pain. In order to improve our understanding of the lumbar spine anatomy and functionality over time, we compared the lumbar vertebrae of Neanderthals with those of anatomically modern humans. The fossil record reports on only two Neanderthal skeletons (i.e., Kebara 2 and Shanidar 3, both predating the appearance of modern humans) with full preservation of the entire lumbar spine. Examination of these early hominids showed that they display natural lumbar kyphosis, with only mild degenerative changes of the lumbar spine (ages at death: 30-35 years, Kebara 2; and 35-50 years, Shanidar 3). This finding is highly unexpected since Neanderthals are known to have had extraordinary physical activity due to demanding living conditions. The adult lumbar spines discussed here therefore show no correlation between high physical activity and degenerative spine disease as known from recent times. We speculate that both the kyphosis itself and the massive and heavily muscled skeleton of Neanderthals are causative for the minimal bone degeneration. We conclude that a kyphotic lumbar spine is the natural anatomy in these two Neanderthal individuals. Future research will reveal if this holds true for the entire Neanderthal species.

High-throughput RNA sequencing reveals structural differences of orthologous brain-expressed genes between western lowland gorillas and humans.

The human brain and human cognitive abilities are strikingly different from those of other great apes despite relatively modest genome sequence divergence. However, little is presently known about the interspecies divergence in gene structure and transcription that might contribute to these phenotypic differences. To date, most comparative studies of gene structure in the brain have examined humans, chimpanzees, and macaque monkeys. To add to this body of knowledge, we analyze here the brain transcriptome of the western lowland gorilla (Gorilla gorilla gorilla), an African great ape species that is phylogenetically closely related to humans, but with a brain that is approximately one-third the size. Manual transcriptome curation from a sample of the planum temporale region of the neocortex revealed 12 protein-coding genes and one noncoding-RNA gene with exons in the gorilla unmatched by public transcriptome data from the orthologous human loci. These interspecies gene structure differences accounted for a total of 134 amino acids in proteins found in the gorilla that were absent from protein products of the orthologous human genes. Proteins varying in structure between human and gorilla were involved in immunity and energy metabolism, suggesting their relevance to phenotypic differences. This gorilla neocortical transcriptome comprises an empirical, not homology- or prediction-driven, resource for orthologous gene comparisons between human and gorilla. These findings provide a unique repository of the sequences and structures of thousands of genes transcribed in the gorilla brain, pointing to candidate genes that may contribute to the traits distinguishing humans from other closely related great apes.

Craniofacial sexual dimorphism patterns and allometry among extant hominids.

Craniofacial sexual dimorphism in primates varies in both magnitude and pattern among species. In the past two decades, there has been an increasing emphasis in exploring the correlations of these patterns with taxonomy and the variation in patterns within and among the craniofacial regions. Scrutinising these relationships for hominids, we decompose the craniofacial morphology in five taxa: Homo sapiens, Pan paniscus, Pan troglodytes, Gorilla gorilla and Pongo pygmaeus. 3D coordinates of 35 traditional landmarks and 61 semilandmarks, covering five ridge curves, are measured for each of 268 adult and sub-adult specimens and analysed using geometric morphometric methods. A multivariate analysis in size-shape space shows that ontogenetic scaling contributes to the development of sexual dimorphism in all five taxa, but to a varying extent. In absolute as well as in relative terms P. pygmaeus shows the greatest allometric component, followed by G. gorilla. Homo is intermediate, while in Pan the non-allometric constituent part contributes a large fraction to the actual sexual dimorphism, most markedly in the pygmy chimpanzee. An eigendecomposition of the five vectors of sexual dimorphism reveals two dimensions independent of allometry. One separates orang-utan sexual dimorphism from the African apes and Homo, and the other differentiates between the great apes and Homo with Pan mediating. We discuss these patterns and speculate on their use as characters for taxonomic analysis in the fossil record.

Development of the palatal size in Pan troglodytes, Hominids and Homo sapiens.

As the hard palate plays an important role in speech production it was the aim of this study whether similarities or dissimilarities in palatal size may allow conclusions about the ability to produce speech in the extant investigated species. The palatal size of Pan troglodytes, Homo sapiens, Australopithecus afarensis, Australopithecus africanus, Australopithecus robustus, Australopithecus boisei, Homo erectus, Homo neanderthalensis and Cro-Magnon has been investigated using euclidian distance matrix analysis (EDMA) and thin-plate-spline analysis. The results show that the palatal size of all australopithecine specimens and H. erectus is very similar to that of P toglodytes, whereas the palatal size of H. neanderthalensis more closely resembles that of H. sapiens. Postnatal development of palatal size in P troglodytes is different from that of H. sapiens. In P troglodytes not only the size of the palate changes but also the form. In H. sapiens there is little change in form, but a continuos uniform growth from infantile to adult specimens. From the results we conclude that in all australopithecine samples which have been investigated, the palatal size is similar to that of P troglodytes. Therefore, it is unlikely that austraopithecine individuals were capable of producing vowels and consonants. The palatal size of H. neandethalensis and Cro-Magnon is similar to that of H. sapiens which may indicate the possibility that they were capable of speech production.

A species' Odyssey: evolution of obstetrical mechanics from Australopithecus Lucy to nowadays.

OBJECTIVE: Study of obstetrical mechanics of Australopithecus Lucy, Homo neanderthalensis and Homo erectus relative to modern Homo sapiens and the Catarrhines. STUDY DESIGN: The material comprised a total of 360 pelves: 3 fossil pelves reconstructed using casts (Australopithecus afarensis Lucy or AL 288-1, Homo erectus KNM-WT 15000, H. neanderthalensis or Kebara 2), 305 female modern adult pelves and 52 female Catarrhine pelves (29 gorillas, 18 chimpanzees, 5 orang-utans). All these pelves were reconstructed in order to carry out 11 pelvimetric measurements. Each measurement was carried out twice and by two different operators. RESULTS: The pelvis of Lucy was platypelloid at each pelvic plane. The pelvic inlet of H. neanderthalensis was anteroposteriorly oval whereas the midplane and the outlet were transversely oval. The pelvis of H. erectus was globally round. In modern women, the inlet was transversely oval. The pelvic midplane and outlet were anteroposteriorly oval. In the great apes, the shape of all three pelvic planes was anteroposteriorly oval. The discriminating value of the various pelvimetry measurements place Australopithecus Lucy, H. neanderthalensis Kebara 2, and H. erectus KNM-WT 15000 close to modern humans and less similar to the great apes. CONCLUSION: Obstetrical mechanics evolved from dystocic delivery with a transverse orientation in Australopithecus to delivery with a modern human-like rotational birth and an increase in the anteroposterior diameters in H. erectus, H. neanderthalensis and modern H. sapiens.

Neuropil distribution in the cerebral cortex differs between humans and chimpanzees.

Increased connectivity of high-order association regions in the neocortex has been proposed as a defining feature of human brain evolution. At present, however, there are limited comparative data to examine this claim fully. We tested the hypothesis that the distribution of neuropil across areas of the neocortex of humans differs from that of one of our closest living relatives, the common chimpanzee. The neuropil provides a proxy measure of total connectivity within a local region because it is composed mostly of dendrites, axons, and synapses. Using image analysis techniques, we quantified the neuropil fraction from both hemispheres in six cytoarchitectonically defined regions including frontopolar cortex (area 10), Broca's area (area 45), frontoinsular cortex (area FI), primary motor cortex (area 4), primary auditory cortex (area 41/42), and the planum temporale (area 22). Our results demonstrate that humans exhibit a unique distribution of neuropil in the neocortex compared to chimpanzees. In particular, the human frontopolar cortex and the frontoinsular cortex had a significantly higher neuropil fraction than the other areas. In chimpanzees these prefrontal regions did not display significantly more neuropil, but the primary auditory cortex had a lower neuropil fraction than other areas. Our results support the conclusion that enhanced connectivity in the prefrontal cortex accompanied the evolution of the human brain. These species differences in neuropil distribution may offer insight into the neural basis of human cognition, reflecting enhancement of the integrative capacity of the prefrontal cortex.

Morfología y morfometría comparada entre los huesecillos auditivos del cerdo común (Sus scrofa domestica) y el Humano (Homo sapiens sapiens) / Comparative study of the morphology between the ossicles of the common pig (Sus scrofa domestica) and human (Homo sapiens sapiens)

El cerdo doméstico (Sus scrofa domestica) ha sido utilizado como modelo para estudiar métodos de implante en el oído medio previo a estudios clínicos humanos. Diferentes investigaciones han destacado la utilidad en este ámbito, como también su alta comparabilidad morfológica en relación con los humanos. Sin embargo, las descripciones anatómicas sobre sus huesecillos auditivos son insuficientes, al igual que las comparaciones en relación con el humano. Por ello, con el objetivo de realizar una descripción detallada de la anatomía de dichos huesecillos comparados con los del humano, se procedió a disecar seis cabezas de cerdo, de las cuales se extrajeron sus huesecillos para observar su morfología. Los resultados preliminares demostraron que la cadena de huesecillos del Cerdo comparte la presencia de las mismas formaciones anatómicas que se pueden identificar en los huesecillos del hombre, pero aún así existen diferencias descriptivas y morfométricas en la morfología de ellas. Los resultados permitieron concluir que existe alta comparabilidad morfológica entre ambas cadenas de huesecillos debido a sus diferencias y similitudes, lo cual lo hace ser un buen modelo didáctico para el estudio y la enseñanza de la morfología auditiva en distintos niveles educacionales.

The domestic Pig (Sus scrofa domestica)has been used as a model to study implants methods in clinical cases of the human middle ear. Different studies have highlighted the usefulness in this area, as well as its high morphological comparability with regard to humans. However, the anatomical descriptions about its ear bones are scarce, as comparisons in relation to the human. Therefore, in order to make a detailed description of the anatomy of these bones compared to human, it was necessary to dissect six pig heads of which its ossicles were removed to observe its morphology. Preliminary results showed that the pig'sossicles share the same anatomical formations that can be identified in the human ones, but there are some descriptive and morphometric differences in its morphology. The results concluded that there is high comparability between both morphological ossicular chains due to their differences and similarities, which makes it a great teaching model for the study and teaching of auditory morphology at different educational levels.