ORCID Profile
0000-0002-1881-2854
Current Organisation
University of the Witwatersrand
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Publisher: Wiley
Date: 11-12-2017
DOI: 10.1002/AR.23738
Abstract: The brain of the crocodile is known to gain in mass allometrically throughout life, and the addition of neurons (as well as non-neurons) appears to play a significant role in this increasing brain mass. We used immunohistochemistry in the brains of 12 Nile crocodiles ranging between 350 g and 86 kg in body mass and 1.99 g to 7.9 g in brain mass to identify the regions of the brain in which neurons immunopositive for doublecortin (DCX), a marker for potential adult neurogenesis, are found. Similar to other reptiles, potential newly born neurons, those immunopositive for DCX, were found throughout the telencephalon, the main and accessory olfactory bulbs and the olfactory tract, and in the cerebellar cortex however, no DCX immunopositive neurons were observed in the diencephalon or brainstem. An apparent moderate decrease in the density of DCX labeled neurons in the olfactory bulbs and tract as well as the cerebellar cortex was observed with increasing brain mass, but the observed qualitative density of labeled neurons within the telencephalon was maintained irrespective of brain mass. Three potential neurogenic zones, within the sulci of the lateral ventricle, were identified, and these are similar to those seen in other reptiles. This study indicates that at least part of the gain in brain mass with age in the Nile crocodile may be accounted for by the potential addition and integration of new neurons into the existing circuitry, especially so for the olfactory system, telencephalon and cerebellar cortex. Anat Rec, 301:659-672, 2018. © 2017 Wiley Periodicals, Inc.
Publisher: Wiley
Date: 04-01-2019
DOI: 10.1002/CNE.24619
Abstract: The diencephalon (dorsal thalamus, ventral thalamus, and epithalamus) and the hypothalamus, play central roles in the processing of the majority of neural information within the central nervous system. Given the interactions of the diencephalon and hypothalamus with virtually all portions of the central nervous system, the comparative analysis of these regions lend key insights into potential neural, evolutionary, and behavioral specializations in different species. Here, we continue our analysis of the brain of the tree pangolin by providing a comprehensive description of the organization of the diencephalon and hypothalamus using a range of standard and immunohistochemical staining methods. In general, the diencephalon and hypothalamus of the tree pangolin follow the organization typically observed across mammals. No unusual structural configurations of the ventral thalamus, epithalamus, or hypothalamus were noted. Within the dorsal thalamus, the vast majority of typically identified nuclear groups and component nuclei were observed. The visual portion of the tree pangolin dorsal thalamus appears to be organized in a manner not dissimilar to that seen in most nonprimate and noncarnivore mammals, and lacks certain features that are present in the closely related carnivores. Within the ventral medial geniculate nucleus, a modular organization, revealed with parvalbumin neuropil immunostaining, is suggestive of specialized auditory processing in the tree pangolin. In addition, a potential absence of hypothalamic cholinergic neurons is suggestive of unusual patterns of sleep. These observations are discussed in an evolutionary and functional framework regarding the phylogeny and life history of the pangolins.
Publisher: Wiley
Date: 09-10-2019
DOI: 10.1002/AR.24265
Abstract: The current study provides a detailed qualitative description of the organization of the cholinergic, catecholaminergic, serotonergic, orexinergic, and GABAergic sleep‐related systems in the brain of the blue wildebeest ( Connocheates taurinus ), along with a quantitative analysis of the pontine cholinergic and noradrenergic neurons, and the hypothalamic orexinergic neurons. The aim of this study was to compare the nuclear organization of these systems to other mammalian species and specifically that reported for other Cetartiodactyla. In the brain of the blue wildebeest, from the basal forebrain to the pons, the nuclear organization of the cholinergic, catecholaminergic, serotonergic, and orexinergic systems, for the most part, showed a corresponding nuclear organization to that reported in other mammals and more specifically the Cetartiodactyla. Furthermore, the description and distribution of the GABAergic system, which was examined through immunostaining for the calcium binding proteins calbindin, calretinin, and parvalbumin, was also similar to that seen in other mammals. These findings indicate that sleep in the blue wildebeest is likely to show typically mammalian features in terms of the global brain activity of the generally recognized sleep states of mammals, but Cetartiodactyl‐specific features of the orexinergic system may act to lower overall daily total sleep time in relation to similar sized non‐Cetartiodactyl mammals. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:1977–1997, 2020. © 2019 American Association for Anatomy
Publisher: Wiley
Date: 22-10-2021
DOI: 10.1002/AR.24788
Abstract: Using tyrosine hydroxylase immunohistochemistry, we describe the nuclear parcellation of the catecholaminergic system in the brains of a lar gibbon ( Hylobates lar ) and a chimpanzee ( Pan troglodytes ). The parcellation of catecholaminergic nuclei in the brains of both apes is virtually identical to that observed in humans and shows very strong similarities to that observed in mammals more generally, particularly other primates. Specific variations of this system in the apes studied include an unusual high‐density cluster of A10dc neurons, an enlarged retrorubral nucleus (A8), and an expanded distribution of the neurons forming the dorsolateral ision of the locus coeruleus (A4). The additional A10dc neurons may improve dopaminergic modulation of the extended amygdala, the enlarged A8 nucleus may be related to the increased use of communicative facial expressions in the hominoids compared to other primates, while the expansion of the A4 nucleus appears to be related to accelerated evolution of the cerebellum in the hominoids compared to other primates. In addition, we report the presence of a compact ision of the locus coeruleus proper (A6c), as seen in other primates, that is not present in other mammals apart from megachiropteran bats. The presence of this nucleus in primates and megachiropteran bats may reflect homology or homoplasy, depending on the evolutionary scenario adopted. The fact that the complement of homologous catecholaminergic nuclei is mostly consistent across mammals, including primates, is advantageous for the selection of model animals for the study of specific dysfunctions of the catecholaminergic system in humans.
Publisher: Wiley
Date: 04-01-2019
DOI: 10.1002/CNE.24620
Abstract: Employing a range of standard and immunohistochemical stains we provide a description of the hippoc al formation in the brain of the tree pangolin. For the most part, the architecture, chemical neuroanatomy, and topological relationships of the component parts of the hippoc al formation of the tree pangolin were consistent with that observed in other mammalian species. Within the hippoc us proper fields CA1, 3, and 4 could be identified with certainty, while CA2 was tentatively identified as a small transitional zone between the CA1 and CA3 fields. Within the dentate gyrus evidence for adult hippoc al neurogenesis at a rate comparable to other mammals was observed. The subicular complex and entorhinal cortex also exhibited isions typically observed in other mammalian species. In contrast to many other mammals, an architecturally and neurochemically distinct CA4 field was observed, supporting Lorente de Nó's proposed CA4 field, at least in some mammalian species. In addition, up to seven laminae were evident in the dentate gyrus. Calretinin immunostaining revealed the three sublamina of the molecular layer, while immunostaining for vesicular glutamate transporter 2 and neurofilament H indicate that the granule cell layer was composed of two sublamina. The similarities and differences observed in the tree pangolin indicate that the hippoc al formation is an anatomically and neurochemically conserved neural unit in mammalian evolution, but minor changes may relate to specific life history features and habits of species.
Publisher: Wiley
Date: 11-05-2017
DOI: 10.1002/CNE.24222
Abstract: Here, we describe the superficial appearance of the brain of the rarely studied tree pangolin. Phylogenetic analyses have placed the pangolins, order Pholidota, as a sister group to the order Carnivora. The majority of features visible on the surface of the tree pangolin brain, and its overall appearance can be described as typically mammalian. The pattern of sulci and gyri, while simple, appears very similar to that observed in carnivores. Two derived features of the Pholidota were observed, the first being the rostral decussation of the pyramidal tract, which instead of occurring at the spinomedullary junction, decussates at the level of the caudal pole of the facial nerve nucleus in the rostral medulla oblongata. This appears to be related to the need for voluntary control of the tongue, with a potentially enlarged corticobulbar tract ending in the hypoglossal nucleus. The second derived feature is the very short spinal cord, which terminates midway along the thoracic vertebrae before giving rise to a long and extensive cauda equina. This foreshortened spinal cord appears to be related to anisotropic growth of the somatic and neural elements following early development of the central nervous system. The olfactory system appears to be generally enlarged and is likely the predominant sense used in foraging. Vision and hearing do not appear specialized based on the relative size of the superior and inferior colliculi, but potential somatic specializations indicate that the somatosensory system is heavily relied upon for food consumption and prehensile tail usage.
Publisher: Wiley
Date: 05-10-2020
DOI: 10.1002/CNE.25036
Abstract: This study provides an analysis of the cytoarchitecture, myeloarchitecture, and chemoarchitecture of the diencephalon (dorsal thalamus, ventral thalamus, and epithalamus) of the banded mongoose ( Mungos mungo ) and domestic ferret ( Mustela putorius furo ). Using architectural and immunohistochemical stains, we observe that the nuclear organization of the diencephalon is very similar in the two species, and similar to that reported in other carnivores, such as the domestic cat and dog. The same complement of putatively homologous nuclei were identified in both species, with only one variance, that being the presence of the perireticular nucleus in the domestic ferret, that was not observed in the banded mongoose. The chemoarchitecture was also mostly consistent between species, although there were a number of minor variations across a range of nuclei in the density of structures expressing the calcium‐binding proteins parvalbumin, calbindin, and calretinin. Thus, despite almost 53 million years since these two species of carnivores shared a common ancestor, strong phylogenetic constraints appear to limit the potential for adaptive evolutionary plasticity within the carnivore order. Apart from the presence of the perireticular nucleus, the most notable difference between the species studied was the physical inversion of the dorsal lateral geniculate nucleus, as well as the lateral posterior and pulvinar nuclei in the domestic ferret compared to the banded mongoose and other carnivores, although this inversion appears to be a feature of the Mustelidae family. While no functional sequelae are suggested, this inversion is likely to result from the altricial birth of Mustelidae species.
Publisher: Wiley
Date: 10-11-2022
DOI: 10.1002/CNE.25431
Abstract: 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.
Publisher: Springer Science and Business Media LLC
Date: 28-04-2023
DOI: 10.1007/S00360-023-01493-6
Abstract: Rest is a state of adaptive inactivity that increases the efficiency of activity by regulating its timing and reducing energy use when activity is not beneficial. Thus, animals can go without rest when specific demands, such as mating, favour being awake. Sexually active male blue wildebeest (bulls) are typically territorial, and it has been reported that when a bull is protecting a harem during the mating season (rut), he neither eats nor rests. We examined the daily activity and inactivity patterns of dominant bulls by means of actigraphy for 3 months, which included the rut. We also measured faecal androgen metabolite (fAM) levels and subcutaneous temperature, both of which have variances known to delineate the rut. During the rut, wildebeest bulls experienced higher levels of activity, fAM, and a greater daily range of subcutaneous temperature. Despite previous reports, the male blue wildebeest rested daily during the rut, and while the amount of rest was low, it was not substantially lower than prior to the rut. The amount of time spent inactive increased substantially after the rut. The timing of daily activity and inactivity patterns did not vary substantially across the recording period. Across the recording period, the average daily ambient temperatures decreased (seasonality), and the subcutaneous temperature followed this pattern, although it was not as marked. It appears that in the post-rut period a substantive increase in time spent at rest occurs, potentially allowing the wildebeest bulls time to recover following a period of intense activity.
Publisher: Wiley
Date: 16-06-2022
DOI: 10.1002/CNE.25353
Abstract: The current study provides a detailed architectural analysis of the subpallial telencephalon of the tree pangolin. In the tree pangolin, the subpallial telencephalon was ided into septal and striatopallidal regions. The septal region contained the septal nuclear complex, diagonal band of Broca, and the bed nuclei of the stria terminalis. The striatopallidal region comprised of the dorsal (caudate, putamen, internal and external globus pallidus) and ventral (nucleus accumbens, olfactory tubercle, ventral pallidum, nucleus basalis, basal part of the substantia innominata, lateral stripe of the striatum, navicular nucleus, and the major island of Calleja) striatopallidal complexes. In the tree pangolin, the organization and numbers of nuclei forming these regions and complexes, their topographical relationships to each other, and the cyto‐, myelo‐, and chemoarchitecture, were found to be very similar to that observed in commonly studied mammals. Minor variations, such as less nuclear parcellation in the bed nuclei of the stria terminalis, may represent species‐specific variations, or may be the result of the limited range of stains used. Given the overall similarity across mammalian species, it appears that the subpallial telencephalon of the mammalian brain is highly conserved in terms of evolutionary changes detectable with the methods used. It is also likely that the functions associated with these nuclei in other mammals can be translated directly to the tree pangolin, albeit with the understanding that the stimuli that produce activity within these regions may be specific to the life history requirements of the tree pangolin.
Publisher: Wiley
Date: 29-06-2022
DOI: 10.1002/CNE.25350
Abstract: The spinal cord of the tree pangolin is known to be very short compared to the overall length of the body and tail. Here, we provide a description of the tree pangolin spinal cord to determine whether the short length contributes to specific structural, and potentially functional, differences. The short spinal cord of the adult tree pangolin, at around 13 cm, terminates at the midthoracic level. Within this shortened spinal cord, we could identify six regions, which from rostral to caudal include the prebrachial, brachial, interramal, crural, postcrural, and caudal regions, with both the brachial and crural regions showing distinct swellings. The chemoarchitecture of coronal sections through these regions confirmed regional assignation, being most readily delineated by the presence of cholinergic neurons forming the intermediolateral column in the interramal region and the sacral parasympathetic nucleus in the postcrural region. The 10 laminae of Rexed were observed throughout the spinal cord and presented with an anatomical organization similar to that observed in other mammals. Despite the shortened length of the tree pangolin spinal cord, the regional and laminar anatomical organization is very similar to that observed in other mammals. This indicates that the functional aspects of the short tree pangolin spinal cord can be inferred from what is known in other mammals.
Publisher: Wiley
Date: 05-10-2018
DOI: 10.1002/AR.23905
Abstract: Relatively little neuroscience research has been focused on artiodactyls. Recent observations of complex social interactions in domestic and wild species suggest that analyses of artiodactyl brain anatomy would be of comparative value. In this study, we examined how the distribution of cortical neuropil space (a proxy for connectivity) varies across representative members of this erse clade. Using image analysis techniques, we quantified the neuropil space in the anterior cingulate cortex (ACC) and the occipital (putative primary visual) cortex (OC) of 12 artiodactyl species from adult specimens. Additionally, we conducted a preliminary investigation of variation in ACC neuropil space in a developmental series of five white-tailed deer (Odocoileus virginianus). Results indicate a consistent pattern of greater neuropil space in the ACC in comparison to the OC among all species, and a gradual increase in ACC neuropil space toward maturity in the white-tailed deer. Given the taxa that have the greatest cortical neuropil space, we hypothesize that such enhanced connectivity might be needed to support behaviors such as group foraging and attentiveness to conspecifics. These results help advance a broader understanding of ersity in neural circuitry in artiodactyls and point to the need for more in-depth comparisons of cortical neuron morphology and organization in this relatively understudied taxonomic group. Anat Rec, 301:1871-1881, 2018. © 2018 Wiley Periodicals, Inc.
Publisher: Wiley
Date: 25-01-2019
DOI: 10.1002/CNE.24605
Publisher: Wiley
Date: 12-10-2021
DOI: 10.1002/AR.24795
Abstract: In the current study, we detail, through the analysis of immunohistochemically stained sections, the morphology and nuclear parcellation of the serotonergic neurons present in the brainstem of a lar gibbon and a chimpanzee. In general, the neuronal morphology and nuclear organization of the serotonergic system in the brains of these two species of apes follow that observed in a range of Eutherian mammals and are specifically very similar to that observed in other species of primates. In both of the apes studied, the serotonergic nuclei could be readily ided into two distinct groups, a rostral and a caudal cluster, which are found from the level of the decussation of the superior cerebellar peduncle to the spinomedullary junction. The rostral cluster is comprised of the caudal linear, supralemniscal, and median raphe nuclei, as well as the six isions of the dorsal raphe nuclear complex. The caudal cluster contains several distinct nuclei and nuclear sub isions, including the raphe magnus nucleus and associated rostral and caudal ventrolateral (CVL) serotonergic groups, the raphe pallidus, and raphe obscurus nuclei. The one deviation in organization observed in comparison to other primate species is an expansion of both the number and distribution of neurons belonging to the lateral ision of the dorsal raphe nucleus in the chimpanzee. It is unclear whether this expansion occurs in humans, thus at present, this expansion sets the chimpanzee apart from other primates studied to date.
Publisher: Wiley
Date: 06-05-2020
DOI: 10.1002/CNE.24922
Publisher: Wiley
Date: 13-06-2019
DOI: 10.1002/CNE.24721
Abstract: The brainstem (midbrain, pons, and medulla oblongata) and cerebellum (diencephalic prosomere 1 through to rhombomere 11) play central roles in the processing of sensorimotor information, autonomic activity, levels of awareness and the control of functions external to the conscious cognitive world of mammals. As such, comparative analyses of these structures, especially the understanding of specializations or reductions of structures with functions that have been elucidated in commonly studied mammalian species, can provide crucial information for our understanding of the behavior of less commonly studied species, like pangolins. In the broadest sense, the nuclear complexes and sub isions of nuclear complexes, the topographical arrangement, the neuronal chemistry, and fiber pathways of the tree pangolin conform to that typically observed across more commonly studied mammalian species. Despite this, variations in regions associated with the locus coeruleus complex, auditory system, and motor, neuromodulatory and autonomic systems involved in feeding, were observed in the current study. While we have previously detailed the unusual locus coeruleus complex of the tree pangolin, the superior olivary nuclear complex of the auditory system, while not exhibiting additional nuclei or having an altered organization, this nuclear complex, particularly the lateral superior olivary nucleus and nucleus of the trapezoid body, shows architectonic refinement. The cephalic decussation of the pyramidal tract, an enlarged hypoglossal nucleus, an additional sub ision of the serotonergic raphe obscurus nucleus, and the expansion of the superior salivatory nucleus, all indicate neuronal specializations related to the myrmecophagous diet of the pangolins.
Publisher: Wiley
Date: 23-07-2021
DOI: 10.1002/CNE.25216
Abstract: 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 s le 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.
Publisher: Wiley
Date: 20-07-2020
DOI: 10.1002/CNE.24972
Publisher: Wiley
Date: 06-03-2020
DOI: 10.1002/CNE.24895
Publisher: Wiley
Date: 04-03-2020
DOI: 10.1002/CNE.24894
Publisher: Wiley
Date: 14-04-2019
DOI: 10.1002/CNE.24692
Publisher: Elsevier BV
Date: 12-2020
Publisher: Wiley
Date: 03-12-2021
DOI: 10.1002/AR.24844
Abstract: Using choline acetyltransferase immunohistochemistry, we describe the nuclear parcellation of the cholinergic system in the brains of two apes, a lar gibbon ( Hylobates lar ) and a chimpanzee ( Pan troglodytes ). The cholinergic nuclei observed in both apes studied are virtually identical to that observed in humans and show very strong similarity to the cholinergic nuclei observed in other primates and mammals more generally. One specific difference between humans and the two apes studied is that, with the specific choline acetyltransferase antibody used, the cholinergic pyramidal neurons observed in human cerebral cortex were not labeled. When comparing the two apes studied and humans to other primates, the presence of a greatly expanded cholinergic medullary tegmental field, and the presence of cholinergic neurons in the intermediate and dorsal horns of the cervical spinal cord are notable variations of the distribution of cholinergic neurons in apes compared to other primates. These neurons may play an important role in the modulation of ascending and descending neural transmissions through the spinal cord and caudal medulla, potentially related to the differing modes of locomotion in apes compared to other primates. Our observations also indicate that the average soma volume of the neurons forming the laterodorsal tegmental nucleus (LDT) is larger than those of the pedunculopontine nucleus (PPT) in both the lar gibbon and chimpanzee. This variability in soma volume appears to be related to the size of the adult derivatives of the alar and basal plate across mammalian species.
Publisher: Wiley
Date: 16-10-2018
DOI: 10.1002/CNE.24519
Abstract: Here, we used a range of immunohistochemical stains, focussing on tyrosine hydroxylase and dopamine-β-hydroxylase, to show that within the pons of tree pangolins clusters of noradrenergic neurons are present. No noradrenergic neurons were observed in the pontine periventricular gray matter (A6 and A4 groups missing), with all noradrenergic neurons being found within the pontine tegmentum (A7 and A5 groups). The tree pangolin is unique in lacking the locus coeruleus (A6) cell group observed in all vertebrates previously studied however, noradrenergic axons and terminal networks were found throughout the cerebral cortex. We propose this is achieved through a unique structural reorganization of this system. First, the number of noradrenergic neurons in the compact portion of the subcoeruleus (A7sc) of the tree pangolin is increased, providing a total number of noradrenergic neurons in the pontine tegmentum (A7diffuse, A7sc, A5) that is equivalent to the entire locus coeruleus complex in related species of similar brain mass. Second, the most medially located noradrenergic neurons of the A7sc have dendrites that extend into the ventrolateral periventricular gray matter, in the location where the A6 neurons should have been located, forming a "pseudo A6" region. Third, the topological relationships of this "pseudo A6" region to other neurochemical systems that interact with the A6 neurons, such as the orexinergic, cholinergic, and serotonergic systems, appear to be maintained. Thus, a unique structural plasticity of this region appears to maintain the standard functions of the locus coeruleus complex in this unusual mammalian species.
Publisher: Wiley
Date: 02-09-2020
DOI: 10.1002/CNE.24999
Publisher: Wiley
Date: 13-09-2020
DOI: 10.1002/CNE.25007
Publisher: Wiley
Date: 21-01-2019
DOI: 10.1002/CNE.24632
Abstract: The present study describes the ipsilateral and contralateral corticocortical and corticothalamic connectivity of the temporal visual areas 20a and 20b in the ferret using standard anatomical tract-tracing methods. The two temporal visual areas are strongly interconnected, but area 20a is primarily connected to the occipital visual areas, whereas area 20b maintains more widespread connections with the occipital, parietal and suprasylvian visual areas and the secondary auditory cortex. The callosal connectivity, although homotopic, consists mainly of very weak anterograde labeling which was more widespread in area 20a than area 20b. Although areas 20a and 20b are well connected with the visual dorsal thalamus, the injection into area 20a resulted in more anterograde label, whereas more retrograde label was observed in the visual thalamus following the injection into area 20b. Most interestingly, comparisons to previous connectional studies of cat areas 20a and 20b reveal a common pattern of connectivity of the temporal visual cortex in carnivores, where the posterior parietal cortex and the central temporal region (PMLS) provide network points required for dorsal and ventral stream interaction enroute to integration in the prefrontal cortex. This pattern of network connectivity is not dissimilar to that observed in primates, which highlights the ferret as a useful animal model to understand visual sensory integration between the dorsal and ventral streams. The data generated will also contribute to a connectomics database, to facilitate cross species analysis of brain connectomes and wiring principles of the brain.
Publisher: Springer Science and Business Media LLC
Date: 03-02-2020
Publisher: Wiley
Date: 13-10-2022
DOI: 10.1002/AR.25092
Abstract: The current study provides an analysis of the cholinergic, catecholaminergic, serotonergic, and orexinergic neuronal populations, or nuclei, in the brain of the lesser hedgehog tenrec, as revealed with immunohistochemical techniques. For all four of these neuromodulatory systems, the nuclear organization was very similar to that observed in other Afrotherian species and is broadly similar to that observed in other mammals. The cholinergic system shows the most variation, with the lesser hedgehog tenrec exhibiting palely immunopositive cholinergic neurons in the ventral portion of the lateral septal nucleus, and the possible absence of cholinergic neurons in the parabigeminal nucleus and the medullary tegmental field. The nuclear complement of the catecholaminergic, serotonergic and orexinergic systems showed no specific variances in the lesser hedgehog tenrec when compared to other Afrotherians, or broadly with other mammals. A striking feature of the lesser hedgehog tenrec brain is a significant mesencephalic flexure that is observed in most members of the Tenrecoidea, as well as the closely related Chrysochlorinae (golden moles), but is not present in the greater otter shrew, a species of the Potomogalidae lineage currently incorporated into the Tenrecoidea. In addition, the cholinergic neurons of the ventral portion of the lateral septal nucleus are observed in the golden moles, but not in the greater otter shrew. This indicates that either complex parallel evolution of these features occurred in the Tenrecoidea and Chrysochlorinae lineages, or that the placement of the Potomogalidae within the Tenrecoidea needs to be re‐examined.
Publisher: Wiley
Date: 25-09-2018
DOI: 10.1002/CNE.24486
Abstract: All domesticated mammals exhibit marked reductions in overall brain size, however, it is unknown whether the corpus callosum (CC), an integral white matter fiber pathway for interhemispheric cortical communication, is affected by domestication differentially or strictly in coordination with changes in brain size. To answer this question, we used quantitative magnetic resonance imaging to compare the midsagittal cross-sectional areas of the CC in 35 carnivore species, including eight wild canids and 13 domestic dogs. We segmented rostro-caudal regions of interest for the CC and evaluated correlations with brain mass. The results of this study indicate that under the influence of domestication in canids, the CC scales to brain size in an allometric relationship that is similar to that of wild canids and other carnivores, with relatively high correlation coefficients observed for all regions, except the rostrum. These results indicate that architectural and energetic considerations are likely to tightly constrain variation in caudal components of the CC relative to overall brain size, however fibers passing through the rostrum, putatively connecting prefrontal cortex, are less constrained and therefore may contribute more toward species-specific differences in connectivity. Given the species ersity of the Canidae and the resurgence of interest in the brain of the domestic dog, further studies aimed at characterizing the neural architecture in domesticated species is likely to provide new insights into the effects of domestication, or artificial selection, on the brain.
Publisher: Springer Science and Business Media LLC
Date: 31-10-2023
Publisher: Wiley
Date: 13-06-2019
DOI: 10.1002/CNE.24709
Abstract: The Nubian ibex (Capra nubiana) occurs in information-rich visual habitats including the edges of cliffs and escarpments. In addition to needing enhanced spatial resolution to find food and detect predators, enhanced visual s ling of the lower visual field would be advantageous for the control of locomotion in such precarious terrains. Using retinal wholemounts and stereology, we sought to measure how the ganglion cell density varies across the retina of the Nubian ibex to reveal which portions of its surroundings are s led with high resolution. We estimated a total of ~1 million ganglion cells in the Nubian ibex retinal ganglion cell layer. Topographic variations of ganglion cell density reveal a temporal area, a horizontal streak, and a dorsotemporal extension, which are topographic retinal features also found in other artiodactyls. In contrast to savannah-dwelling artiodactyls, the horizontal streak of the Nubian ibex appears loosely organized possibly reflecting a reduced predation risk in mountainous habitats. Estimates of spatial resolving power (~17 cycles/degree) for the temporal area would be reasonable to facilitate foraging in the frontal visual field. Embedded in the dorsotemporal extension, we also found an unusual dorsotemporal area not yet reported in any other mammal. Given its location and spatial resolving power (~6 cycles/degree), this specialization enhances visual s ling toward the lower visual field, which would be advantageous for visually guided locomotion. This study expands our understanding of the retinal organization in artiodactyls and offers insights on the importance of vision for the Nubian ibex ecology.
Publisher: Wiley
Date: 21-02-2018
DOI: 10.1002/JMOR.20804
Abstract: Anomodontia was the most successful herbivorous clade of the mammalian stem lineage (non-mammalian synapsids) during the late Permian and Early Triassic. Among anomodonts, Dicynodontia stands apart because of the presence of an osseous beak that shows evidence of the insertion of a cornified sheath, the r hotheca. In this study, fourteen anomodont specimens were microCT-scanned and their trigeminal canals reconstructed digitally to understand the origin and evolution of trigeminal nerve innervation of the r hotheca. We show that the pattern of innervation of the anomodont "beak" is more similar to that in chelonians (the nasopalatine branch is enlarged and innervates the premaxillary part of the r hotheca) than in birds (where the nasopalatine and maxillary branches play minor roles). The nasopalatine branch is noticeably enlarged in the beak-less basal anomodont Patranomodon, suggesting that this could be an anomodont or chainosaur synapomorphy. Our analyses suggest that the presence or absence of tusks and postcanine teeth are often accompanied by corresponding variations of the rami innervating the caniniform process and the alveolar region, respectively. The degree of ossification of the canal for the nasal ramus of the ophthalmic branch also appears to correlate with the presence of a nasal boss. The nasopalatine canal is absent from the premaxilla in the Bidentalia as they uniquely show a large plexus formed by the internal nasal branch of the maxillary canal instead. The elongated shape of this plexus in Lystrosaurus supports the hypothesis that the rostrum evolved as an elongation of the subnarial region of the snout. Finally, the atrophied and variable aspect of the trigeminal canals in Myosaurus supports the hypothesis that this genus had a reduced upper r hotheca.
Publisher: Wiley
Date: 19-08-2020
DOI: 10.1002/CNE.24989
Publisher: Wiley
Date: 16-12-2019
DOI: 10.1002/CNE.24823
Abstract: 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 s les 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.
Publisher: Wiley
Date: 26-11-2017
DOI: 10.1002/CNE.24349
Abstract: Gigantopyramidal neurons, referred to as Betz cells in primates, are characterized by large somata and extensive basilar dendrites. Although there have been morphological descriptions and drawings of gigantopyramidal neurons in a limited number of species, quantitative investigations have typically been limited to measures of soma size. The current study thus employed two separate analytical approaches: a morphological investigation using the Golgi technique to provide qualitative and quantitative somatodendritic measures of gigantopyramidal neurons across 19 mammalian species from 7 orders and unbiased stereology to compare the soma volume of layer V pyramidal and gigantopyramidal neurons in primary motor cortex between 11 carnivore and 9 primate species. Of the 617 neurons traced in the morphological analysis, 181 were gigantopyramidal neurons, with deep (primarily layer V) pyramidal (n = 203) and superficial (primarily layer III) pyramidal (n = 233) neurons quantified for comparative purposes. Qualitatively, dendritic morphology varied considerably across species, with some (sub)orders (e.g., artiodactyls, perissodactyls, feliforms) exhibiting bifurcating, V-shaped apical dendrites. Basilar dendrites exhibited idiosyncratic geometry across and within taxonomic groups. Quantitatively, most dendritic measures were significantly greater in gigantopyramidal neurons than in superficial and deep pyramidal neurons. Cluster analyses revealed that most taxonomic groups could be discriminated based on somatodendritic morphology for both superficial and gigantopyramidal neurons. Finally, in agreement with Brodmann, gigantopyramidal neurons in both the morphological and stereological analyses were larger in feliforms (especially in the Panthera species) than in other (sub)orders, possibly due to specializations in muscle fiber composition and musculoskeletal systems.
Publisher: Wiley
Date: 30-10-2018
DOI: 10.1002/CNE.24545
Publisher: Wiley
Date: 08-2020
DOI: 10.1002/CNE.24985
Publisher: Wiley
Date: 23-09-2021
DOI: 10.1002/AR.24775
Abstract: Employing orexin‐A immunohistochemical staining we describe the nuclear parcellation of orexinergic neurons in the hypothalami of a lar gibbon and a chimpanzee. The clustering of orexinergic neurons within the hypothalamus and the terminal networks follow the patterns generally observed in other mammals, including laboratory rodents, strepsirrhine primates and humans. The orexinergic neurons were found within three distinct clusters in the ape hypothalamus, which include the main cluster, zona incerta cluster and optic tract cluster. In addition, the orexinergic neurons of the optic tract cluster appear to extend to a more rostral and medial location than observed in other species, being observed in the tuberal region in the anterior ventromedial aspect of the hypothalamus. While orexinergic terminal networks were observed throughout the brain, high density terminal networks were observed within the hypothalamus, medial and intralaminar nuclei of the dorsal thalamus, and within the serotonergic and noradrenergic regions of the midbrain and pons, which is typical for mammals. The expanded distribution of orexinergic neurons into the tuberal region of the ape hypothalamus, is a feature that needs to be investigated in other primate species, but appears to correlate with orexin gene expression in the same region of the human hypothalamus, but these neurons are not revealed with immunohistochemical staining in humans. Thus, it appears that apes have a broader distribution of orexinergic neurons compared to other primate species, but that the neurons within this extension of the optic tract cluster in humans, while expressing the orexin gene, do not produce the neuropeptide.
Publisher: Wiley
Date: 18-10-2018
DOI: 10.1002/CNE.24510
Abstract: This study employed a range of neuroanatomical stains to determine the organization of the main and accessory olfactory systems within the brain of the tree pangolin. The tree pangolin has a typically mammalian olfactory system, but minor variations were observed. The main olfactory system is comprised of the layered main olfactory bulb (MOB), the anterior olfactory nucleus (AON), the rostral olfactory cortex (including the taenia tecta, anterior hippoc al continuation and induseum griseum), the olfactory tubercle (Tu), the lateral olfactory tract (lot) and the olfactory limb of the anterior commissure, the nucleus of the lateral olfactory tract (NLOT), the piriform cortex (PIR) and a typically mammalian rostral migratory stream (RMS). The accessory olfactory system included the layered accessory olfactory bulb (AOB) and the nucleus of the accessory olfactory tract (NAOT). Volumetric analysis of the relative size of the MOB and PIR indicate that the tree pangolin has an olfactory system that occupies a proportion of the brain typical for the majority of mammals. Within the MOB, the glomeruli of the tree pangolin, at 200 μm diameter, are larger than observed in most other mammalian species, and the MOB lacks a distinct internal plexiform layer. In addition, the laminate appearance of the NLOT was not observed in the tree pangolin. The accessory olfactory system appears to lack the posterior compartment of the accessory olfactory bulb. These observations are contextualized in relation to olfactory-mediated behaviors in pangolins.
Publisher: Wiley
Date: 25-01-2019
DOI: 10.1002/CNE.24631
Abstract: The present study describes the ipsilateral and contralateral corticocortical and corticothalamic connectivity of the occipital visual areas 17, 18, 19, and 21 in the ferret using standard anatomical tract-tracing methods. In line with previous studies of mammalian visual cortex connectivity, substantially more anterograde and retrograde label was present in the hemisphere ipsilateral to the injection site compared to the contralateral hemisphere. Ipsilateral reciprocal connectivity was the strongest within the occipital visual areas, while weaker connectivity strength was observed in the temporal, suprasylvian, and parietal visual areas. Callosal connectivity tended to be strongest in the homotopic cortical areas, and revealed a similar areal distribution to that observed in the ipsilateral hemisphere, although often less widespread across cortical areas. Ipsilateral reciprocal connectivity was observed throughout the visual nuclei of the dorsal thalamus, with no contralateral connections to the visual thalamus being observed. The current study, along with previous studies of connectivity in the cat, identified the posteromedial lateral suprasylvian visual area (PMLS) as a distinct network hub external to the occipital visual areas in carnivores, implicating PMLS as a potential gateway to the parietal cortex for dorsal stream processing. These data will also contribute to a macro connectome database of the ferret brain, providing essential data for connectomics analyses and cross-species analyses of connectomes and brain connectivity matrices, as well as providing data relevant to additional studies of cortical connectivity across mammals and the evolution of cortical connectivity variation.
Publisher: Wiley
Date: 22-01-2019
DOI: 10.1002/CNE.24630
Abstract: The present study describes the ipsilateral and contralateral cortico-cortical and cortico-thalamic connectivity of the parietal visual areas, posterior parietal caudal cortical area (PPc) and posterior parietal rostral cortical area (PPr), in the ferret using standard anatomical tract-tracing methods. The two isions of posterior parietal cortex of the ferret are strongly interconnected, however area PPc shows stronger connectivity with the occipital and suprasylvian visual cortex, while area PPr shows stronger connectivity with the somatomotor cortex, reflecting the functional specificity of these two areas. This pattern of connectivity is mirrored in the contralateral callosal connections. In addition, PPc and PPr are connected with the visual and somatomotor nuclei of the dorsal thalamus. Numerous connectional similarities exist between the posterior parietal cortex of the ferret (PPc and PPr) and the cat (area 7 and 5), indicative of the homology of these areas within the Carnivora. These findings highlight the existence of a frontoparietal network as a shared feature of the organization of parietal cortex across Euarchontoglires and Laurasiatherians, with the degree of expression varying in relation to the expansion and areal complexity of the posterior parietal cortex. This observation indicates that the ferret is a potentially valuable experimental model animal for understanding the evolution and function of the posterior parietal cortex and the frontoparietal network across mammals. The data generated will also contribute to a connectomics database, to further cross-species analyses of connectomes and illuminate wiring principles of cortical connectivity across mammals.
Publisher: Wiley
Date: 07-2021
DOI: 10.1002/CNE.25202
Abstract: We examined the number, distribution, and immunoreactivity of the infracortical white matter neuronal population, also termed white matter interstitial cells (WMICs), throughout the telencephalic white matter of an adult female chimpanzee. Staining for neuronal nuclear marker (NeuN) revealed WMICs throughout the infracortical white matter, these cells being most numerous and dense close to the inner border of cortical layer VI, decreasing significantly in density with depth in the white matter. Stereological analysis of NeuN‐immunopositive cells revealed an estimate of approximately 137.2 million WMICs within the infracortical white matter of the chimpanzee brain studied. Immunostaining revealed subpopulations of WMICs containing neuronal nitric oxide synthase (nNOS, approximately 14.4 million in number), calretinin (CR, approximately 16.7 million), very few WMICs containing parvalbumin (PV), and no calbindin‐immunopositive neurons. The nNOS, CR, and PV immunopositive WMICs, possibly all inhibitory neurons, represent approximately 22.6% of the total WMIC population. As the white matter is affected in many cognitive conditions, such as schizophrenia, autism, epilepsy, and also in neurodegenerative diseases, understanding these neurons across species is important for the translation of findings of neural dysfunction in animal models to humans. Furthermore, studies of WMICs in species such as apes provide a crucial phylogenetic context for understanding the evolution of these cell types in the human brain.
Publisher: Wiley
Date: 14-10-2020
DOI: 10.1002/CNE.25047
Abstract: Employing cyto‐, myelo‐, and chemoarchitectural staining techniques, we analyzed the structure of the hippoc al formation in the banded mongoose and domestic ferret, species belonging to the two carnivoran superfamilies, which have had independent evolutionary trajectories for the past 55 million years. Our observations indicate that, despite the time since sharing a last common ancestor, these species show extensive similarities. The four major portions of the hippoc al formation (cornu Ammonis, dentate gyrus, subicular complex, and entorhinal cortex) were readily observed, contained the same internal sub isions, and maintained the topological relationships of these sub isions that could be considered typically mammalian. In addition, adult hippoc al neurogenesis was observed in both species, occurring at a rate similar to that observed in other mammals. Despite the overall similarities, several differences to each other, and to other mammalian species, were observed. We could not find evidence for the presence of the CA2 and CA4 fields of the cornu Ammonis region. In the banded mongoose the dentate gyrus appears to be comprised of up to seven lamina, through the sublamination of the molecular and granule cell layers, which is not observed in the domestic ferret. In addition, numerous subtle variations in chemoarchitecture between the two species were observed. These differences may contribute to an overall variation in the functionality of the hippoc al formation between the species, and in comparison to other mammalian species. These similarities and variations are important to understanding to what extent phylogenetic affinities and constraints affect potential adaptive evolutionary plasticity of the hippoc al formation.
Publisher: Wiley
Date: 20-10-2020
DOI: 10.1002/CNE.25046
Abstract: The current study provides an analysis of the cytoarchitecture, myeloarchitecture, and chemoarchitecture of the amygdaloid body of the banded mongoose ( Mungos mungo ) and domestic ferret ( Mustela putorius furo ). Using architectural and immunohistochemical stains, we observe that the organization of the nuclear and cortical portions of the amygdaloid complex is very similar in both species. The one major difference is the presence of a cortex‐amygdala transition zone observed in the domestic ferret that is absent in the banded mongoose. In addition, the chemoarchitecture is, for the most part, quite similar in the two species, but several variances, such as differing densities of neurons expressing the calcium‐binding proteins in specific nuclei are noted. Despite this, certain aspects of the chemoarchitecture, such as the cholinergic innervation of the magnocellular ision of the basal nuclear cluster and the presence of doublecortin expressing neurons in the shell ision of the accessory basal nuclear cluster, appear to be consistent features of the Eutherian mammal amygdala. The domestic ferret presented with an overall lower myelin density throughout the amygdaloid body than the banded mongoose, a feature that may reflect artificial selection in the process of domestication for increased juvenile‐like behavior in the adult domestic ferret, such as a muted fear response. The shared, but temporally distant, ancestry of the banded mongoose and domestic ferret allows us to generate observations relevant to understanding the relative influence that phylogenetic constraints, adaptive evolutionary plasticity, and the domestication process may play in the organization and chemoarchitecture of the amygdaloid body.
Publisher: Wiley
Date: 18-09-2020
DOI: 10.1002/CNE.25000
Publisher: American Association for the Advancement of Science (AAAS)
Date: 30-04-2021
Abstract: An in-depth look at mammalian brain size evolution prompts a reevaluation of a traditional paradigm.
Publisher: Academy of Science of South Africa
Date: 29-09-2021
Location: United States of America
No related grants have been discovered for Paul Manger.