Neuropil Variation in the Prefrontal, Motor, and Visual Cortex of Six Felids.

INTRODUCTION: Felids have evolved a specialized suite of morphological adaptations for obligate carnivory. Although the musculoskeletal anatomy of the Felidae has been studied extensively, the comparative neuroanatomy of felids is relatively unexplored. Little is known about how variation in the cerebral anatomy of felids relates to species-specific differences in sociality, hunting strategy, or activity patterns. METHODS: We quantitatively analyzed neuropil variation in the prefrontal, primary motor, and primary visual cortices of six species of Felidae (Panthera leo, Panthera uncia, Panthera tigris, Panthera leopardus, Acinonyx jubatus, Felis sylvestris domesticus) to investigate relationships with brain size, neuronal cell parameters, and select behavioral and ecological factors. Neuropil is the dense, intricate network of axons, dendrites, and synapses in the brain, playing a critical role in information processing and communication between neurons. RESULTS: There were significant species and regional differences in neuropil proportions, with African lion, cheetah, and tiger having more neuropil in all three cortical regions in comparison to the other species. Based on regression analyses, we find that the increased neuropil fraction in the prefrontal cortex supports social and behavioral flexibility, while in the primary motor cortex, this facilitates the neural activity needed for hunting movements. Greater neuropil fraction in the primary visual cortex may contribute to visual requirements associated with diel activity patterns. CONCLUSION: These results provide a cross-species comparison of neuropil fraction variation in the Felidae, particularly the understudied Panthera, and provide evidence for convergence of the neuroanatomy of Panthera and cheetahs.

Reproducibility of leftward planum temporale asymmetries in two genetically isolated populations of chimpanzees (Pan troglodytes).

Once considered a hallmark of human uniqueness, brain asymmetry has emerged as a feature shared with several other species, including chimpanzees, one of our closest living relatives. Most notable has been the discovery of asymmetries in homologues of cortical language areas in apes, particularly in the planum temporale (PT), considered a central node of the human language network. Several lines of evidence indicate a role for genetic mechanisms in the emergence of PT asymmetry; however, the genetic determinants of cerebral asymmetries have remained elusive. Studies in humans suggest that there is heritability of brain asymmetries of the PT, but this has not been explored to any extent in chimpanzees. Furthermore, the potential influence of non-genetic factors has raised questions about the reproducibility of earlier observations of PT asymmetry reported in chimpanzees. As such, the present study was aimed at examining both the heritability of phenotypic asymmetries in PT morphology, as well as their reproducibility. Using magnetic resonance imaging, we evaluated morphological asymmetries of PT surface area (mm2) and mean depth (mm) in captive chimpanzees (n = 291) derived from two genetically isolated populations. Our results confirm that chimpanzees exhibit a significant population-level leftward asymmetry for PT surface area, as well as significant heritability in the surface area and mean depth of the PT. These results conclusively demonstrate the existence of a leftward bias in PT asymmetry in chimpanzees and suggest that genetic mechanisms play a key role in the emergence of anatomical asymmetry in this region.

Hippocampal neurogenesis in the C57BL/6J mice at early adulthood following prenatal alcohol exposure.

We examined the effect of chronic prenatal alcohol exposure (PAE) on the process of adult neurogenesis in C57BL/6J mice at early adulthood (PND 56). Pregnant mice, and their in utero litters, were exposed to alcohol, through oral gavage, on gestational days 7-16, with recorded blood alcohol concentrations averaging 184 mg/dL (CA group). Two control groups, sucrose (CAc) and non-treated (NTc) control groups were also examined. The brains of pups at PND 56 from each experimental group were sectioned in a sagittal plane, and stained for Nissl substance with cresyl violet, and immunostained for Ki-67 which labels proliferative cells and doublecortin (DCX) for immature neurons. Morphologically, the neurogenic pattern was identical in all three groups studied. Populations of Ki-67 immunopositive cells in the dentate gyrus were not statistically significantly different between the experimental groups and there were no differences between the sexes. Thus, the PAE in this study does not appear to have a strong effect on the proliferative process in the adult hippocampus. In contrast, the numbers of immature neurons, labeled with DCX, was statistically significantly lower in the prenatal alcohol exposed mice compared with the two control groups. Alcohol significantly lowered the number of DCX hippocampal cells in the male mice, but not in the female mice. This indicates that the PAE appears to lower the rate of conversion of proliferative cells to immature neurons and this effect of alcohol is sexually dimorphic. This lowered number of immature neurons in the hippocampus appears to mirror hippocampal dysfunctions observed in FASD children.

Nuclear parcellation and numbers of orexinergic neurons in five species of larger brained birds.

The orexinergic/hypocretinergic system, while having several roles, appears to be a key link in the balance between arousal and food intake. In birds, to date, this system has only been examined anatomically in four species, all with brains smaller than 3.5 g and of limited phylogenetic range. Here, using orexin-A immunohistochemistry, we describe the distribution, morphology, and nuclear parcellation of orexinergic neurons within the hypothalami of a Congo gray and a Timneh gray parrot, a pied crow, an emu, and a common ostrich. These birds represent a broad phylogeny, with brains ranging in size from 7.85 to 26.5 g. Within the hypothalami of the species studied, the orexinergic neurons were organized in two clusters, and a densely packed paraventricular hypothalamic nucleus cluster located within the medial hypothalamus (Hyp), but not contacting the ventricle, and a more loosely packed lateral hypothalamic cluster in the lateral Hyp. Stereological analysis revealed a strong correlation, using phylogenetic generalized least squares regression analyses, between brain mass and the total number of orexinergic neurons, as well as soma parameters such as volume and area. Orexinergic axonal terminals evinced two types of boutons, larger and the smaller en passant boutons. Unlike the orexinergic system in mammals, which has several variances in cluster organization, that of the birds studied, in the present and previous studies, currently shows organizational invariance, despite the differences in brain and body mass, phylogenetic relationships, and life-histories of the species studied.

Comparative anatomy of the caudate nucleus in canids and felids: Associations with brain size, curvature, cross-sectional properties, and behavioral ecology.

The evolutionary history of canids and felids is marked by a deep time separation that has uniquely shaped their behavior and phenotype toward refined predatory abilities. The caudate nucleus is a subcortical brain structure associated with both motor control and cognitive, emotional, and executive functions. We used a combination of three-dimensional imaging, allometric scaling, and structural analyses to compare the size and shape characteristics of the caudate nucleus. The sample consisted of MRI scan data obtained from six canid species (Canis lupus lupus, Canis latrans, Chrysocyon brachyurus, Lycaon pictus, Vulpes vulpes, Vulpes zerda), two canid subspecies (Canis lupus familiaris, Canis lupus dingo), as well as three felids (Panthera tigris, Panthera uncia, Felis silvestris catus). Results revealed marked conservation in the scaling and shape attributes of the caudate nucleus across species, with only slight deviations. We hypothesize that observed differences in caudate nucleus size and structure for the domestic canids are reflective of enhanced cognitive and emotional pathways that possibly emerged during domestication.

The evolutions of large brain size in mammals: the 'over-700-gram club quartet'.

The current paper details our developing understanding of the evolution of large brains in mammals. In order to do this, we first define brains that we consider to be large--those that have passed the apparent 700-gram ceiling on brain mass evolution in the class Mammalia. The over-700-gram club includes certain species within the genus Homo, order Cetacea, order Proboscidea, and suborder Pinnipedia. Our analysis suggests that selection for body size appears to be the most important factor in the evolution of large brain size, but there also appear to be internal morphophysiological constraints on large brain size evolution that need to be overcome in order for brains to break the 700-gram barrier. These two aspects appear to be common themes in the evolution of large brains. This significantly diminishes the explanatory value of selection for greater cognitive capacities as a principal factor in the evolution of enlarged brain sizes above the 700-gram threshold.

Qualitative and quantitative aspects of the microanatomy of the African elephant cerebellar cortex.

The current study provides a number of novel observations on the organization and structure of the cerebellar cortex of the African elephant by using a combination of basic neuroanatomical and immunohistochemical stains with Golgi and stereologic analysis. While the majority of our observations indicate that the cerebellar cortex of the African elephant is comparable to other mammalian species, several features were unique to the elephant. The three-layered organization of the cerebellar cortex, the neuronal types and some aspects of the expression of calcium-binding proteins were common to a broad range of mammalian species. The Lugaro neurons observed in the elephant were greatly enlarged in comparison to those of other large-brained mammals, suggesting a possible alteration in the processing of neural information in the elephant cerebellar cortex. Analysis of Golgi impregnations indicated that the dendritic complexity of the different interneuron types was higher in elephants than other mammals. Expression of parvalbumin in the parallel fibers and calbindin expressed in the stellate and basket cells also suggested changes in the elephant cerebellar neuronal circuitry. The stereologic analysis confirmed and extended previous observations by demonstrating that neuronal density is low in the elephant cerebellar cortex, providing for a larger volume fraction of the neuropil. With previous results indicating that the elephants have the largest relative cerebellar size amongst mammals, and one of the absolutely largest mammalian cerebella, the current observations suggest that the elephants have a greater volume of a potentially more complexly organized cerebellar cortex compared to other mammals. This quantitatively larger and more complex cerebellar cortex likely represents part of the neural machinery required to control the complex motor patterns involved in movement of the trunk and the production of infrasonic vocalizations.

Resequencing of the TMF-1 (TATA Element Modulatory Factor) regulated protein (TRNP1) gene in domestic and wild canids.

BACKGROUND: Cortical folding is related to the functional organization of the brain. The TMF-1 regulated protein (TRNP1) regulates the expansion and folding of the mammalian cerebral cortex, a process that may have been accelerated by the domestication of dogs. The objectives of this study were to sequence the TRNP1 gene in dogs and related canid species, provide evidence of its expression in dog brain and compare the genetic variation within dogs and across the Canidae. The gene was located in silico to dog chromosome 2. The sequence was experimentally confirmed by amplifying and sequencing the TRNP1 exonic and promoter regions in 72 canids (36 purebred dogs, 20 Gy wolves and wolf-dog hybrids, 10 coyotes, 5 red foxes and 1 Gy fox). RESULTS: A partial TRNP1 transcript was isolated from several regions in the dog brain. Thirty genetic polymorphisms were found in the Canis sp. with 17 common to both dogs and wolves, and only one unique to dogs. Seven polymorphisms were observed only in coyotes. An additional 9 variants were seen in red foxes. Dogs were the least genetically diverse. Several polymorphisms in the promoter and 3'untranslated region were predicted to alter TRNP1 function by interfering with the binding of transcriptional repressors and miRNAs expressed in neural precursors. A c.259_264 deletion variant that encodes a polyalanine expansion was polymorphic in all species studied except for dogs. A stretch of 15 nucleotides that is found in other mammalian sequences (corresponding to 5 amino acids located between Pro58 and Ala59 in the putative dog protein) was absent from the TRNP1 sequences of all 5 canid species sequenced. Both of these aforementioned coding sequence variations were predicted to affect the formation of alpha helices in the disordered region of the TRNP1 protein. CONCLUSIONS: Potentially functionally important polymorphisms in the TRNP1 gene are found within and across various Canis species as well as the red fox, and unique differences in protein structure have evolved and been conserved in the Canidae compared to all other mammalian species.

The folded shape of mammalian brains allows the cerebral cortex and other structures to attain a large surface area to fit into the skull. Over the past several thousand years, the domestication of dogs has led to increased folding as well as a great variety of skull size and shape amongst various breeds. Cortical folding, also known as gyrification, is regulated by numerous genes, including one that encodes the TMF-1 regulated protein (TRNP1). The TRNP1 protein in the brain affects the development of specialized cells in the brain that are involved in gyrification. It is not known whether a functionally distinct TRNP1 protein in dogs is responsible for the increased folding. Variations in the DNA sequence of the gene that encodes TRNP1 may be responsible for these dramatic changes in brain structure in dogs. This study sought to discover the differences in the TRNP1 DNA sequence in seventy-two canids, represented by thirty-six dogs of various breeds, twenty gray wolves and wolf-dog hybrids, ten coyotes, five red foxes and one gray fox.After finding evidence of the expression of this gene in dog brain, we located thirty genetic changes or variants in the canids, with seventeen common to both dogs and wolves, and only one unique to dogs. Another seven of these genetic variants were observed only in coyotes. An additional nine variants were seen in red foxes. Dogs were the least genetically diverse species, an expected result of the inbreeding that characterizes domestication. Several of these changes may affect the function of the TRNP1 gene by affecting the binding of other biomolecules to regions in the DNA which regulate this gene. This study also found two other changes, one only found in dogs, and the other one only found in canids (compared to all other mammalian TRNP1 proteins) may change the length and three-dimensional structure and hence the function of the TRNP1 protein. This study concluded that numerous, potentially functionally significant dissimilarities in the TRNP1 gene exist between dogs and their wild relatives, as well as between canid and all other mammalian species.

Orexinergic neurons in the hypothalami of an Asiatic lion, an African lion, and a Southeast African cheetah.

Employing orexin-A immunohistochemistry, we describe the distribution, morphology, and nuclear parcellation of orexinergic neurons within the hypothalami of an Asiatic lion (Panthera leo subsp. persica), an African lion (Panthera leo subsp. melanochaita), and a Southeast African cheetah (Acinonyx jubatus subsp. jubatus). In all three felids, the clustering of large, bipolar, and multipolar hypothalamic orexinergic neurons primarily follows the pattern observed in other mammals. The orexinergic neurons were found, primarily, to form three distinct clusters-the main, zona incerta, and optic tract clusters. In addition, large orexinergic neurons were observed in the ventromedial supraoptic region of the hypothalamus, where they are not typically observed in other species. As has been observed in cetartiodactyls and the African elephant, a cluster of small, multipolar orexinergic neurons, the parvocellular cluster, was observed in the medial zone of the hypothalamus in all three felids, although this parvocellular cluster has not been reported in other carnivores. In both subspecies of lions, but not the cheetah, potential orexin-immunopositive neurons were observed in the paraventricular hypothalamic nucleus, supraoptic nucleus, the lateral part of the retrochiasmatic area, and the inner layer of the median eminence. The distribution and parcellation of orexinergic neurons in the hypothalami of the three felids studied appear to be more complex than observed in many other mammals and for the two subspecies of lion may be even more complex. These findings are discussed in terms of potential technical concerns, phylogenetic variations of this system, and potentially associated functional aspects of the orexinergic system.

Sex estimation from dimensions of the base of the skull in Black South Africans.

The pelvis and the skull are the two most utilised skeletal elements to estimate sex from skeletonised remains due to their sexually dimorphic traits. However, as increasingly more fragmented remains have been presented for analyses, other bones and their fragments have now been subjected to analyses for sex estimation. In the skull particularly, the base has shown to survive harsh conditions. In this study the foramen magnum region was explored in Black South Africans to estimate sex during forensic analyses. Seven measurements of the foramen magnum and surrounding areas were measured in 120 male and female crania and subjected to discriminant function analyses. The average accuracies for the stepwise discriminant functions ranged from 60-71% whilst the average accuracies for the direct discriminant functions ranged from 63-69%. The average accuracies obtained in this study are similar to other studies performed using the foramen magnum. However, these average accuracies are much lower than other skeletal elements that have been used for sex estimation in South Africans. Thus, the equations in this study should be used with caution and only in the absence of more accurate elements. The cranial base has always shown to have a low to moderate expression of sexual dimorphism. The cranial base of Black South Africans is no different.

Broca's area homologue in chimpanzees (Pan troglodytes): probabilistic mapping, asymmetry, and comparison to humans.

Neural changes that occurred during human evolution to support language are poorly understood. As a basis of comparison to humans, we used design-based stereological methods to estimate volumes, total neuron numbers, and neuron densities in Brodmann's areas 44 and 45 in both cerebral hemispheres of 12 chimpanzees (Pan troglodytes), one of our species' closest living relatives. We found that the degree of interindividual variation in the topographic location and quantitative cytoarchitecture of areas 44 and 45 in chimpanzees was comparable to that seen in humans from previous studies. However, in contrast to the documented asymmetries in humans, we did not find significant population-level hemispheric asymmetry for any measures of areas 44 and 45 in chimpanzees. Furthermore, there was no relationship between asymmetries of stereological data and magnetic resonance imaging-based measures of inferior frontal gyrus morphology or hand preference on 2 different behavioral tasks. These findings suggest that Broca's area in the left hemisphere expanded in relative size during human evolution, possibly as an adaptation for our species' language abilities.

The distribution, number, and certain neurochemical identities of infracortical white matter neurons in the brains of a southern lesser galago, a black-capped squirrel monkey, and a crested macaque.

In the current study, we examined the number, distribution, and aspects of the neurochemical identities of infracortical white matter neurons, also termed white matter interstitial cells (WMICs), in the brains of a southern lesser galago (Galago moholi), a black-capped squirrel monkey (Saimiri boliviensis boliviensis), and a crested macaque (Macaca nigra). Staining for neuronal nuclear marker (NeuN) revealed WMICs throughout the infracortical white matter, these cells being most dense close to inner cortical border, decreasing in density with depth in the white matter. Stereological analysis of NeuN-immunopositive cells revealed estimates of approximately 1.1, 10.8, and 37.7 million WMICs within the infracortical white matter of the galago, squirrel monkey, and crested macaque, respectively. The total numbers of WMICs form a distinct negative allometric relationship with brain mass and white matter volume when examined in a larger sample of primates where similar measures have been obtained. In all three primates studied, the highest densities of WMICs were in the white matter of the frontal lobe, with the occipital lobe having the lowest. Immunostaining revealed significant subpopulations of WMICs containing neuronal nitric oxide synthase (nNOS) and calretinin, with very few WMICs containing parvalbumin, and none containing calbindin. The nNOS and calretinin immunopositive WMICs represent approximately 21% of the total WMIC population; however, variances in the proportions of these neurochemical phenotypes were noted. Our results indicate that both the squirrel monkey and crested macaque might be informative animal models for the study of WMICs in neurodegenerative and psychiatric disorders in humans.

Amplification of potential thermogenetic mechanisms in cetacean brains compared to artiodactyl brains.

To elucidate factors underlying the evolution of large brains in cetaceans, we examined 16 brains from 14 cetartiodactyl species, with immunohistochemical techniques, for evidence of non-shivering thermogenesis. We show that, in comparison to the 11 artiodactyl brains studied (from 11 species), the 5 cetacean brains (from 3 species), exhibit an expanded expression of uncoupling protein 1 (UCP1, UCPs being mitochondrial inner membrane proteins that dissipate the proton gradient to generate heat) in cortical neurons, immunolocalization of UCP4 within a substantial proportion of glia throughout the brain, and an increased density of noradrenergic axonal boutons (noradrenaline functioning to control concentrations of and activate UCPs). Thus, cetacean brains studied possess multiple characteristics indicative of intensified thermogenetic functionality that can be related to their current and historical obligatory aquatic niche. These findings necessitate reassessment of our concepts regarding the reasons for large brain evolution and associated functional capacities in cetaceans.

Wernicke's area homologue in chimpanzees (Pan troglodytes) and its relation to the appearance of modern human language.

Human language is distinctive compared with the communication systems of other species. Yet, several questions concerning its emergence and evolution remain unresolved. As a means of evaluating the neuroanatomical changes relevant to language that accompanied divergence from the last common ancestor of chimpanzees, bonobos and humans, we defined the cytoarchitectonic boundaries of area Tpt, a component of Wernicke's area, in 12 common chimpanzee brains and used design-based stereologic methods to estimate regional volumes, total neuron number and neuron density. In addition, we created a probabilistic map of the location of area Tpt in a template chimpanzee brain coordinate space. Our results show that chimpanzees display significant population-level leftward asymmetry of area Tpt in terms of neuron number, with volume asymmetry approaching significance. Furthermore, asymmetry in the number of neurons in area Tpt was positively correlated with asymmetry of neuron numbers in Brodmann's area 45, a component of Broca's frontal language region. Our findings support the conclusion that leftward asymmetry of Wernicke's area originated prior to the appearance of modern human language and before our divergence from the last common ancestor. Moreover, this study provides the first evidence of covariance between asymmetry of anterior and posterior cortical regions that in humans are important to language and other higher order cognitive functions.

Inhibitory interneurons of the human prefrontal cortex display conserved evolution of the phenotype and related genes.

Inhibitory interneurons participate in local processing circuits, playing a central role in executive cognitive functions of the prefrontal cortex. Although humans differ from other primates in a number of cognitive domains, it is not currently known whether the interneuron system has changed in the course of primate evolution leading to our species. In this study, we examined the distribution of different interneuron subtypes in the prefrontal cortex of anthropoid primates as revealed by immunohistochemistry against the calcium-binding proteins calbindin, calretinin and parvalbumin. In addition, we tested whether genes involved in the specification, differentiation and migration of interneurons show evidence of positive selection in the evolution of humans. Our findings demonstrate that cellular distributions of interneuron subtypes in human prefrontal cortex are similar to other anthropoid primates and can be explained by general scaling rules. Furthermore, genes underlying interneuron development are highly conserved at the amino acid level in primate evolution. Taken together, these results suggest that the prefrontal cortex in humans retains a similar inhibitory circuitry to that in closely related primates, even though it performs functional operations that are unique to our species. Thus, it is likely that other significant modifications to the connectivity and molecular biology of the prefrontal cortex were overlaid on this conserved interneuron architecture in the course of human evolution.

The brain of the African wild dog. II. The olfactory system.

Employing a range of neuroanatomical stains, we detail the organization of the main and accessory olfactory systems of the African wild dog. The organization of both these systems follows that typically observed in mammals, but variations of interest were noted. Within the main olfactory bulb, the size of the glomeruli, at approximately 350 µm in diameter, are on the larger end of the range observed across mammals. In addition, we estimate that approximately 3,500 glomeruli are present in each main olfactory bulb. This larger main olfactory bulb glomerular size and number of glomeruli indicates that enhanced peripheral processing of a broad range of odorants is occurring in the main olfactory bulb of the African wild dog. Within the accessory olfactory bulb, the glomeruli did not appear distinct, rather forming a homogenous syncytia-like arrangement as seen in the domestic dog. In addition, the laminar organization of the deeper layers of the accessory olfactory bulb was indistinct, perhaps as a consequence of the altered architecture of the glomeruli. This arrangement of glomeruli indicates that rather than parcellating the processing of semiochemicals peripherally, these odorants may be processed in a more nuanced and combinatorial manner in the periphery, allowing for more rapid and precise behavioral responses as required in the highly social group structure observed in the African wild dog. While having a similar organization to that of other mammals, the olfactory system of the African wild dog has certain features that appear to correlate to their environmental niche.

The brain of the African wild dog. IV. The visual system.

The variegated pelage and social complexity of the African wild dog (Lycaon pictus) hint at the possibility of specializations of the visual system. Here, using a range of architectural and immunohistochemical stains, we describe the systems-level organization of the image-forming, nonimage forming, oculomotor, and accessory optic, vision-associated systems in the brain of one representative individual of the African wild dog. For all of these systems, the organization, in terms of location, parcellation and topology (internal and external), is very similar to that reported in other carnivores. The image-forming visual system consists of the superior colliculus, visual dorsal thalamus (dorsal lateral geniculate nucleus, pulvinar and lateral posterior nucleus) and visual cortex (occipital, parietal, suprasylvian, temporal and splenial visual regions). The nonimage forming visual system comprises the suprachiasmatic nucleus, ventral lateral geniculate nucleus, pretectal nuclear complex and the Edinger-Westphal nucleus. The oculomotor system incorporates the oculomotor, trochlear and abducens cranial nerve nuclei as well as the parabigeminal nucleus, while the accessory optic system includes the dorsal, lateral and medial terminal nuclei. The extent of similarity to other carnivores in the systems-level organization of these systems indicates that the manner in which these systems process visual information is likely to be consistent with that found, for example, in the well-studied domestic cat. It would appear that the sociality of the African wild dog is dependent upon the processing of information extracted from the visual system in the higher-order cognitive and affective neural systems.

The brain of the African wild dog. III. The auditory system.

The large external pinnae and extensive vocal repertoire of the African wild dog (Lycaon pictus) has led to the assumption that the auditory system of this unique canid may be specialized. Here, using cytoarchitecture, myeloarchitecture, and a range of immunohistochemical stains, we describe the systems-level anatomy of the auditory system of the African wild dog. We observed the cochlear nuclear complex, superior olivary nuclear complex, lateral lemniscus, inferior colliculus, medial geniculate body, and auditory cortex all being in their expected locations, and exhibiting the standard subdivisions of this system. While located in the ectosylvian gyri, the auditory cortex includes several areas, resembling the parcellation observed in cats and ferrets, although not all of the auditory areas known from these species could be identified in the African wild dog. These observations suggest that, broadly speaking, the systems-level anatomy of the auditory system, and by extension the processing of auditory information, within the brain of the African wild dog closely resembles that observed in other carnivores. Our findings indicate that it is likely that the extraction of the semantic content of the vocalizations of African wild dogs, and the behaviors generated, occurs beyond the classically defined auditory system, in limbic or association neocortical regions involved in cognitive functions. Thus, to obtain a deeper understanding of how auditory stimuli are processed, and how communication is achieved, in the African wild dog compared to other canids, cortical regions beyond the primary sensory areas will need to be examined in detail.

Brain of the African wild dog. I. Anatomy, architecture, and volumetrics.

The African wild dog is endemic to sub-Saharan Africa and belongs to the family Canidae which includes domestic dogs and their closest relatives (i.e., wolves, coyotes, jackals, dingoes, and foxes). The African wild dog is known for its highly social behavior, co-ordinated pack predation, and striking vocal repertoire, but little is known about its brain and whether it differs in any significant way from that of other canids. We employed gross anatomical observation, magnetic resonance imaging, and classical neuroanatomical staining to provide a broad overview of the structure of the African wild dog brain. Our results reveal a mean brain mass of 154.08 g, with an encephalization quotient of 1.73, indicating that the African wild dog has a relatively large brain size. Analysis of the various structures that comprise their brains and their topological inter-relationships, as well as the areas and volumes of the corpus callosum, ventricular system, hippocampus, amygdala, cerebellum and the gyrification index, all reveal that the African wild dog brain is, in general, similar to that of other mammals, and very similar to that of other carnivorans. While at this level of analysis we do not find any striking specializations within the brain of the African wild dog, apart from a relatively large brain size, the observations made indicate that more detailed analyses of specific neural systems, particularly those involved in sensorimotor processing, sociality or cognition, may reveal features that are either unique to this species or shared among the Canidae to the exclusion of other Carnivora.

Comparative neocortical neuromorphology in felids: African lion, African leopard, and cheetah.

The present study examines cortical neuronal morphology in the African lion (Panthera leo leo), African leopard (Panthera pardus pardus), and cheetah (Acinonyx jubatus jubatus). Tissue samples were removed from prefrontal, primary motor, and primary visual cortices and investigated with a Golgi stain and computer-assisted morphometry to provide somatodendritic measures of 652 neurons. Although neurons in the African lion were insufficiently impregnated for accurate quantitative dendritic measurements, descriptions of neuronal morphologies were still possible. Qualitatively, the range of spiny and aspiny neurons across the three species was similar to those observed in other felids, with typical pyramidal neurons being the most prominent neuronal type. Quantitatively, somatodendritic measures of typical pyramidal neurons in the cheetah were generally larger than in the African leopard, despite similar brain sizes. A MARsplines analysis of dendritic measures correctly differentiated 87.4% of complete typical pyramidal neurons between the African leopard and cheetah. In addition, unbiased stereology was used to compare the soma size of typical pyramidal neurons (n = 2,238) across all three cortical regions and gigantopyramidal neurons (n = 1,189) in primary motor and primary visual cortices. Both morphological and stereological analyses indicated that primary motor gigantopyramidal neurons were exceptionally large across all three felids compared to other carnivores, possibly due to specializations related to the felid musculoskeletal systems. The large size of these neurons in the cheetah which, unlike lions and leopards, does not belong to the Panthera genus, suggests that exceptionally enlarged primary motor gigantopyramidal neurons evolved independently in these felid species.