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Postural control in man: the phylogenetic perspective.

Erect posture in man is a recent affordance from an evolutionary perspective. About eight million years ago, the stock from which modern humans derived split off from the ape family, and from around sixty-thousand years ago, modern man developed. Upright gait and manipulations while standing pose intricate cybernetic problems for postural control. The trunk, having an older evolutionary history than the extremities, is innervated by medially descending motor systems and extremity muscles by the more recent, laterally descending systems. Movements obviously require concerted actions from both systems. Research in rats has demonstrated the interdependencies between postural control and the development of fluent walking. Only 15 days after birth, adult-like fluent locomotion emerges and is critically dependent upon postural development. Vesttibular deprivation induces a retardation in postural development and, consequently, a retarded development of adult-like locomotion. The cerebellum obviously has an important role in mutual adjustments in postural control and extremity movements, or, in coupling the phylogenetic older and newer structures. In the human, the cerebellum develops partly after birth and therefore is vulnerable to adverse perinatal influences. Such vulnerability seems to justify focusing our scientific research efforts onto the development of this structure.

Animals↗

Bihemispheric contribution to motor recovery after stroke: A longitudinal study with transcranial doppler ultrasonography.

BACKGROUND: Both cerebral hemispheres seem to contribute to motor recovery after stroke. We studied the effect of motor activity on cerebral blood flow in both hemispheres at different stages of stroke evolution. METHODS: Thirty patients with hemiplegic stroke and 30 controls were included. Patients were examined within the first week (T1), 1 month (T2) and 6 months after stroke (T3). All subjects performed a 2-min sequential thumb-to-finger opposition task while blood flow velocities in both middle cerebral arteries were measured with transcranial Doppler ultrasonography (TCD). RESULTS: Contralateral movement caused a higher increase in blood flow velocity than ipsilateral movement in controls (p < 0.0001). On the healthy side, patients showed a striking increase with ipsilateral movement (affected hand), which was similar to the increase with contralateral movement (normal hand) at all stages. On the damaged side, the increase with contralateral movement (affected hand) was low and was similar to the increase with ipsilateral movement (normal hand) at T1 and T2; however, at T3 the increase with contralateral movement was higher and the pattern of response was similar to that found in controls. CONCLUSIONS: TCD can trace the evolution of brain motor output following stroke. Compensatory activation of the healthy side of the brain may be already present soon after stroke, whereas function of the damaged side may improve during several months.

Aged↗

Social cognition and psychopathology in an evolutionary perspective. Current status and proposals for research.

The phylogenetic and ontogenetic developments of social cognition have been a major research focus of evolutionary and developmental psychology. Theory of mind or so-called Machiavellian intelligence, that is the capacity to infer mental states of other individuals and to manipulate them in order to maximise social success, probably emerged due to the need to cope with an increasingly complex social environment. Studies on social reasoning suggest disturbances of mental state attribution in psychiatric disorders. However, apart from autism spectrum disorders, the systematic evaluation of social cognition is still in its infancy, and the present data are ambiguous due to methodological difficulties. Based on the concept of the modular organisation of the mind, a stepwise investigation of social cognition in psychiatric disorders is proposed, including clinical description and available standardised methods. The specific characteristics of psychiatric disorders in respect of social cognition, therefore, may vary according to the hierarchical organisation of the social module. Systematic studies on social reasoning processes in psychiatric disorders may provide new insights also useful for the development of coping strategies in cognitive-behavioural therapy.

Biological Evolution↗

Convergence of complex cognitive abilities in cetaceans and primates.

What examples of convergence in higher-level complex cognitive characteristics exist in the animal kingdom? In this paper I will provide evidence that convergent intelligence has occurred in two distantly related mammalian taxa. One of these is the order Cetacea (dolphins, whales and porpoises) and the other is our own order Primates, and in particular the suborder anthropoid primates (monkeys, apes, and humans). Despite a deep evolutionary divergence, adaptation to physically dissimilar environments, and very different neuroanatomical organization, some primates and cetaceans show striking convergence in social behavior, artificial 'language' comprehension, and self-recognition ability. Taken together, these findings have important implications for understanding the generality and specificity of those processes that underlie cognition in different species and the nature of the evolution of intelligence.

Animal Communication↗

Cortical orofacial motor representation in Old World monkeys, great apes, and humans. I. Quantitative analysis of cytoarchitecture.

Social life in anthropoid primates is mediated by interindividual communication, involving movements of the orofacial muscles for the production of vocalization and gestural expression. Although phylogenetic diversity has been reported in the auditory and visual communication systems of primates, little is known about the comparative neuroanatomy that subserves orofacial movement. The current study reports results from quantitative image analysis of the region corresponding to orofacial representation of primary motor cortex (Brodmann's area 4) in several catarrhine primate species (Macaca fascicularis, Papio anubis, Pongo pygmaeus, Gorilla gorilla, Pan troglodytes, and Homo sapiens) using the Grey Level Index method. This cortical region has been implicated in the execution of skilled motor activities such as voluntary facial expression and human speech. Density profiles of the laminar distribution of Nissl-stained neuronal somata were acquired from high-resolution images to quantify cytoarchitectural patterns. Despite general similarity in these profiles across catarrhines, multivariate analysis showed that cytoarchitectural patterns of individuals were more similar within-species versus between-species. Compared to Old World monkeys, the orofacial representation of area 4 in great apes and humans was characterized by an increased relative thickness of layer III and overall lower cell volume densities, providing more neuropil space for interconnections. These phylogenetic differences in microstructure might provide an anatomical substrate for the evolution of greater volitional fine motor control of facial expressions in great apes and humans.

Aged↗

A pallial visual area in the telencephalon of the bony fish Polypterus.

The dorsal (P2) and lateral (P3) pallial zones of bichirs receive a substantial projection from nucleus medianus of the posterior tuberculum. Although nucleus medianus does not receive a direct retinal input, its close proximity to ascending tectal efferents suggests that it might receive a tectal input and form a segment of a retinotectal-tubercular pathway to the pallium. In order to test this possibility, evoked responses to light flashes were electrophysiologically recorded and tectal efferents were experimentally determined in bichirs. These experiments suggest that a single visual field exists across the dorsal and lateral pallial zones and that this field is mediated by nucleus medianus, which does receive a direct tectal projection. This visual pathway appears to be uniquely derived and not homologous to any other known pathway in tetrapods. Furthermore, these results support the contention that the P2 and P3 pallial zones in bichirs are subdivisions of a single pallial zone.

Animals↗

Parcellation and area-area connectivity as a function of neocortex size.

Via the accumulation of data from across the neuroanatomy literature, we estimate the manner in which (i) the number of neocortical areas varies with neocortex size, and (ii) the number of area-area connections varies with neocortex size. Concerning parcellation, we find that the number of areas scales approximately as the 1/3 power of gray matter volume, or, equivalently, as the square root of the total number of neocortical neurons. A consequence of this is that the average number of neurons per area also scales approximately as the square root of the total number of areas. Concerning area-area connectivity, we find evidence that the total number of area-area connections scales as the square of the number of areas. These scaling results help constrain theories about the principles underlying neocortical organization.

Animals↗

Afferent and efferent connections of the lateral and medial pallia of the silver lamprey.

Injections of the carbocyanine dye, DiI, into the lateral pallium of the silver lamprey reveal that this pallial region receives bilateral inputs from the olfactory bulbs, dorsomedial telencephalic neuropil, and the habenular nuclei, and ipsilateral inputs from the septum, preoptic area, medial pallium, thalamus, and, possibly, the striatum. The efferent projections of the lateral pallium form dorsal (olfacto-habenular tract of Heier) and ventral (olfacto-thalamic and hypothalamic tracts of Heier) bundles. The dorsal bundle terminates ipsilaterally in the dorsal pallium, medial pallium, habenular nuclei, and pretectum and contralaterally in the habenular nuclei and, possibly, the dorsal pallium. The ventral bundle terminates ipsilaterally in the septum, striatum, and preoptic areas and bilaterally within the hypothalamus. Injections of DiI into the medial pallium reveal bilateral inputs to this pallial formation from the olfactory bulbs, the dorsomedial telencephalic neuropil, septum, habenular nuclei, thalamic nuclei, preoptic area and hypothalamus, as well as ipsilateral inputs from the lateral pallium, dorsal isthmal grey and midbrain tegmentum. The efferent projections of the medial pallium form dorsal, ventral and descending bundles. The dorsal bundle terminates ipsilaterally in the dorsal and lateral pallia and in the olfactory bulb. The ventral bundle terminates ipsilaterally in the dorsal pallium and bilaterally within the lateral pallium and in preoptic and hypothalamic areas. The descending bundle terminates bilaterally in thalamic and hypothalamic areas and in the pretectum and optic tectum. These data support a number of earlier hypotheses concerning pallial homologues in lampreys and other vertebrates but suggest that the earlier hypothesis of an olfactory origin of the telencephalon of craniates should be rejected.

Afferent Pathways↗

Afferent and efferent connections of the central prosencephalic nucleus in the Pacific hagfish.

Injections of horseradish peroxidase, or the carbocyanine dye Dil, into the central prosencephalic nucleus of hagfishes reveal reciprocal connections with the septum, preoptic area, habenula, thalamus and hypothalamus. Efferent projections also occur to the olfactory bulb, pallium, posterior tubercle and tegmentum and to the contralateral central prosencephalic nucleus. These connections suggest that the central prosencephalic nucleus of hagfishes may be homologous to portions of the medial pallium and ventral thalamus of other craniates.

Afferent Pathways↗

Neuroanatomical and histochemical evidence for the presence of common lateral line and inner ear efferents and of efferents to the basilar papilla in a frog, Xenopus laevis.

The claimed absence of efferents to the basilar papilla in frogs was reexamined in Xenopus laevis with the use of acetylcholinesterase (AChE) histochemistry. We also tested the presence of common lateral line and inner ear efferents by employing neuroanatomical tract tracing techniques. Our data show some AChE-positive fibers to the basilar papilla and all other sensory epithelia of the inner ear in larval and postmetamorphic frogs. Labeling of anterior lateral line and inner ear fibers with different fluorescing dextran amines resulted in a few double labeled efferent cells in the brainstem. Examination of the branching patterns of anterior lateral line efferents revealed collaterals to the inner ear, predominantly to the saccule and the lagena. In addition, two animals showed common efferents between the anterior lateral line and the basilar papilla. In derived anurans, such as ranids, which reportedly lack efferents to the basilar papilla, the basilar papillary afferents have a peripheral course that differs from that in salamanders and Xenopus. If such efferents are, indeed, absent, we propose that changed cues for pathway selection may have enabled only the afferents, and not the efferents, to reach the basilar papilla in derived frogs.

Acetylcholinesterase↗

The octavolateralis system and Mauthner cell: interactions and questions.

This paper is an overview of some of the major points to arise in the accompanying contributions of this special symposium issue. The symposium papers arose out of discussions among investigators interested in the inner ear and Mauthner cell, with the focus on hydrodynamic components that activate the Mauthner cell through the octavolateralis system. The intention of the symposium was to investigate the possibility of using our knowledge of the Mauthner system to help understand acoustic processing by the ear, and of using our knowledge of fish hearing to better understand Mauthner cell function. This is the first attempt to take a broad look at both systems to see how they might function together. As such, these proceedings can serve as a mini-tutorial for investigators interested in one system or the other. In this summary paper we also identify some of the major uncertainties in our understanding of the ear-Mauthner connection. These include questions about: (1) the identity of the acoustic stimuli that are neuroethologically relevant to the Mauthner system; (2) the relative importance of the various octavolateralis inputs (acoustic, vestibular, or lateral line); (3) the contribution of the different various acoustic endorgans to the Mauthner system; (4) whether the Mauthner system can distinguish sound source location, and (5) whether Mauthner neurobiology is compatible with the prevailing model (the phase model) for determining sound source location by fishes. We believe these issues provide potentially useful avenues of future investigation that should give important insights in both acoustic processing by fish and the function of the mauthner system.

Animals↗

The limbic system of tetrapods: a comparative analysis of cortical and amygdalar populations.

Recent studies of the limbic system of tetrapods have made data available that challenge some of the long-held tenets of forebrain evolution. Using the basic principle of parsimony--that the best hypotheses concerning homologies are those requiring the fewest number of evolutionary changes--we have reevaluated comparisons of tetrapod limbic systems. Given the current data, the following points appear to be justified: (1) the common ancestors of reptiles and mammals had a well-developed limbic system in which the basic subdivisions and connections of the amygdalar nuclei were established; (2) the ventral part of the lateral pallium in amphibians appears to be a single structure which corresponds to at least four areas in reptiles: centromedial DVR, ventral anterior amygdala, lateral amygdala, and part of the lateral cortex; (3) the medial pallium in amphibians appears to be homologous with the dorsal and medial cortices in reptiles and with the general and hippocampal cortices in mammals: (4) the cortical targets of the main olfactory bulb in reptiles and mammals appear to be homologous, and their common ancestor probably had a corresponding olfactory pallial field; (5) the targets of the accessory olfactory bulb in amphibians, reptiles, and mammals appear to be homologous, with the exception of nucleus sphericus in reptiles, which lacks an obvious homologue in non-reptiles.

Amphibians↗

Comparative aspects of forebrain organization in the ray-finned fishes: touchstones or not?

The comparative interpretation of forebrain organization of the ray-finned fishes presents a number of challenges. The telencephalon develops by an eversion process, and much of its surface is ventricular rather than pial. A topological map of the ventricular surface shows that the area ventralis (presumed subpallium) is bordered dorsally by the dorsomedial (DM) zone of the area dorsalis, which is, in turn, bordered by the olfactory-recipient dorsoposterior (DP) zone of the area dorsalis. Within the diencephalon small dorsal and ventral thalami and a large group of migrated nuclei assigned to the posterior tuberculum (but of uncertain status) are present. Both the dorsal thalamus and the migrated nuclei of the posterior tuberculum project to the telencephalon. A review of the known connections of the four major zones of the area dorsalis together with their topological positions leads to the following tentative interpretations. Zone DP is homologous to the primary olfactory cortex. There is not enough information concerning the dorsodorsal zone (DD) to speculate on its comparative relationships. Zone DM is a limbic-like area lying between the subpallium and the olfactory pallium and is possibly homologous to the pallial amygdala. The dorsolateral zone (DL) displays a pattern of connectivity with the cerebellum and the optic tectum suggesting a possible homology with non-olfactory, non-limbic pallial areas. Caveats are raised concerning the above interpretations of DM and DL. One or both of them may have evolved independently of the telencephalic zones in tetrapods and may thus represent examples of homoplasy.

Animals↗

Evolution of taste and solitary chemoreceptor cell systems.

Vertebrates possess four distinct chemosensory systems distinguishable on the basis of structure, innervation and utilization: olfaction, taste, solitary chemoreceptor cells (SCC) and the common chemical sense (free nerve endings). Of these, taste and the SCC sense rely on secondary receptor cells situated in the epidermis and synapsing on sensory nerve fibers innervating them near their base. The SCC sense occurs in anamniote aquatic craniates, including hagfish, and may be used for feeding or predator avoidance. The sense of taste occurs only in vertebrates and is always utilized for feeding. The SCC system achieves a high degree of specialization in two teleosts: sea robins (Prionotus) and rocklings (Ciliata). In sea robins, SCCs are abundant on the three anterior fin rays of the pectoral fin which are free of fin webbing and are used in active exploration of the substrate. Behavioral and physiological studies show that this SCC system responds to feeding cues and drives feeding behavior. It is connected centrally like a somatosensory system. In contrast, the specialized SCC system of rocklings occurs on the anterior dorsal fin which actively samples the surrounding water. This system responds to mucus substances and may serve as a predator detector. The SCC system in rocklings is connected centrally like a gustatory system. Taste buds contain multiple receptor cell types, including a serotonergic Merkel-like cell. Taste receptor cells respond to nutritionally relevant substances. Due to similarities between SCCs and one type of taste receptor cell, the suggestion is made that taste buds may be compound sensory organs that include some cells related to SCCs and others related to cutaneous Merkel cells. The lack of taste buds in hagfish and their presence in all vertebrates may indicate that the phylogenetic development of taste buds coincided with the elaboration of head structures at the craniate-vertebrate transition.

Animals↗

A lacertilian dorsal retinorecipient thalamus: a re-investigation in the old-world lizard Podarcis hispanica.

The aim of this work is to delineate the retinorecipient cell groups of the dorsal thalamus of lizards and to study some of the differential connections in order to help to understand the evolution of the visual system in tetrapods. Tract-tracing and immunohistochemical techniques were applied to the retinorecipient dorsal thalamus of the lizard Podarcis hispanica. The retina of Podarcis projects to four areas of the dorsal thalamus: nucleus ovalis (Ov), intergeniculate leaflet (IGL), dorsal lateral geniculate nucleus (GLD) and dorsolateral anterior nucleus (DLA). Nucleus ovalis shows a clear cell plate/neuropile organization and projects to the ventral thalamus. Thus, it seems to belong to the ventral rather than to the dorsal thalamus. The IGL contains large cells reactive for GABA and/or NPY immunohistochemistry. It is interconnected with the supra/retrochiasmatic hypothalamus and projects to the opposite thalamus and to the ipsilateral tectum. The caudal DLA, which lacks both GABA- and NPY-like immunoreactive cells, is reached by a few thin retinal fibers, although distal dendrites of DLA cells enter the GLD, suggesting an important retinal input. The DLA projects to the medial and dorsal telencephalic cortices. The GLD is the main retinorecipient thalamic structure that projects to the telencephalon. It shows a crude laminar organization in which cell plate neurons project to the ipsilateral pallial thickening, but it does not receive a descending projection from the visual telencephalon and thus differs from the GLD of other amniotic vertebrates. In the context of present knowledge, these results suggest that an IGL homologue is present in all tetrapods studied, whereas Ov seems to be restricted to diapsid vertebrates. Moreover, our data suggest that a unimodal visual projection to the telencephalon (arising from the GLD) first appeared in reptiles by segregation from a limbic (multimodal) thalamo-telencephalic pathway.

Animals↗

Anti-Zebrin II immunopositivity in the cerebellum and octavolateral nuclei in two species of stingrays.

Anti-Zebrin II is an antibody directed against a 36kDa aldolase epitope expressed by Purkinje cells. Two patterns of Zebrin II immunolabeling have been described in the cerebellar corpus. In mammalian cerebella, the anti-Zebrin II labels longitudinal zones of Purkinje cells, whereas in teleosts, all Purkinje cells of the cerebellar corpus are Zebrin II immunopositive. An outgroup analysis is required to determine which of these distribution patterns represents the primitive condition for jawed vertebrates. The sister group of the Osteichthyans (rayfinned fishes, amphibians, and amniotes) is the Chondrichthyans (sharks, skates, and rays). In the present study the distribution of Zebrin II immunopositivity was examined in the Atlantic stingray and the Southern stingray. Western-blot analysis demonstrates that the Zebrin II antibody recognizes an antigen of the same molecular weight in stingrays, teleosts, and mammals. In stingrays, anti-Zebrin II immunohistochemistry reveals a staining pattern in which all Purkinje cells are immunopositive, no banding pattern or zonal compartmentation is observed. Purkinje cell axon projections to the cerebellar nucleus and the octaval nuclei are also revealed. Within the octaval nuclei, immunopositive Purkinje cell axon terminals and boutons en passant were found in the anterior, descending, and posterior nuclei. These immunopositive profiles are found throughout these nuclei, but they are most dense in the lateral and ventral portions. Except for the dorsolateralmost portion, the magnocellular nucleus does not appear to receive Purkinje cell inputs. Based on these results it is concluded that the Zebrin II distribution pattern in which all Purkinje cells of the cerebellar corpus are immunopositive is the primitive condition for jawed vertebrates.

Animals↗