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The postnatal development of somatosensory callosal connections after partial lesions of somatosensory areas
No full textThe distribution of S1 (first somatosensory area) and S2 (second somatosensory area) neurons projecting to the contralateral S2 was studied with horseradish peroxidase in normal adult cats and in cats aged between 129 and 248 days in which the injected S2 area had been deprived of some of its input by an earlier lesion (on postnatal days 3 to 30; day of birth = day 1) of ipsilateral S1, alone or combined with a lesion of contralateral S2. In animals with S1 lesions, as in the normal controls, labeled neurons were selectively distributed to the regions of the trunk representation and to parts of the forelimb and hindlimb representations; however, the normally acallosal region in the forepaw representation contained scattered labeled neurons in three of the four animals whose S1 had been lesioned during the first postnatal week. In these animals, the distribution of labeled neurons in the contralateral S2 was apparently normal. Furthermore, the additional lesion of this area during the first postnatal week (one animal) did not increase the degree of filling-in of the normally acallosal parts of S1. The partial filling-in of the acallosal parts of S1 is probably due to the preservation to adulthood of some of the callosal neurons which are present in these regions during the early postnatal life. Possibly, these neurons did not disappear (or lose their callosal axons) because the neonatal lesion (i) allowed their successful competition for terminal space in contralateral S2 or (ii) induced a reorganization of the peripheral input to this area
Internal representations of movement in the cerebral cortex as revealed by the analysis of reaching
No full textReaching movements have the well-defined goal of bringing the hand to the location of an object of interest. For neuroscientists a basic problem to be solved is how the nervous system transforms the visual information concerning the location of the object in space into a pattern of muscle activity necessary to bring the hand to it. According to Descartes, spirits passing from the eyes impinge on the pineal gland, causing it to lean in one direction or another; this leaning of the gland pulls on filaments (nerves) attached to the muscles. Modern treatments, instead, tend to decompose this process into sequences of transformations between informational representations. Such transformations lead from a description of the target in visual coordinates to an expression of the movement in muscle space by way of various internal representations. The organization of such internal representations has implications for the types of transformations actually performed in the brain. Recent psychophysical, neurophysiological, and computational approaches to study the cortical representations of reaching movements are yielding complementary data on this issue
Postnatal shaping of callosal connections from sensory areas
No full textHorseradish peroxidase (HRP) was injected unilaterally into the first and second visual areas (V1 and V2; areas 17 and 18) of 20 kittens aged between 2 and 90 days and into the second somatosensory area (S2) of 16 kittens aged between 1 and 52 days. The radial and tangential (normal and parallel to the pial surface, respectively) distributions of neurons giving origin to callosal axons (callosal neurons) were studied. In adult cats, callosal efferent zones (CZs) are defined by the distribution of callosal neurons. CZs occupy, in the visual cortices, tangentially and radially restricted parts of areas 17, 18, 19 of the lateral suprasylvian gyrus and in the somatosensory cortices, parts of S1 and S2. At birth, callosal neurons are distributed throughout the tangential extent of visual and somatosensory areas; they are also more widespread in depth than in the adult. During the first postnatal month, as a result of the gradual disappearance of callosal neurons from parts of the visual and somatosensory areas, the adult CZs emerge. The CZ in areas 17 and 18 undergoes a further tangential reduction during the second and third postnatal months
Optic ataxia as result of the breakdown of the global tuning fields of parietal neurons
No full textOptic ataxia is characterized by an impaired visual control of the direction of arm reaching to a visual target, accompanied by defective hand orientation and grip formation. In humans, optic ataxia is associated with lesions of the superior parietal lobule (SPL), which also affect visually guided saccades and other forms of eye-hand coordination. In the last 10 years, anatomical and physiological studies of the SPL have shed new light on the role of parietal cortex in the control of combined eye-hand movements to visual targets, and on the underlying distributed network which links parietal to frontal cortex. A main emerging functional feature of SPL neurones seems to be their capacity to combine, in a spatially congruent fashion, different directional eye- and hand-related information, that any coding scheme so far proposed, considers essential for the composition of motor commands for reaching. This integration occurs within the global tuning field of parietal neurones, is context-dependent and involves eye and hand information that shares the same directional properties. Depending on task demands, this integration of signals can result in the representation of different reference frames for coordinated eye-hand movements. The dynamic operations occurring within the global tuning fields might depend, at least in part, on the reciprocal sets of association connections linking the SPL and the premotor areas of the frontal lobe. From this picture, the SPL emerges as both a main source of visual input to the frontal cortex and a key structure for visuomotor integration based on re-entrant signalling and, therefore, as a crucial node in the visual control of movement. It is hypothesized that in parietal patients, the directional errors that characterize reaching are a consequence of the breakdown of the combination of directional eye and hand information within the global tuning fields of parietal neurones. In these patients, the spatial match among information about target location, eye and hand position, and movement direction would be prevented, so as to impair the composition of visually guided eye-hand movements. This breakdown could be dependent, at least in part, on the failure of a re-entrant frontoparietal signalling, an obvious consequence of the degeneration of the cortico-cortical systems linking parietal and frontal cortex. Cortico-cortical connections are, in fact, essential for shaping the dynamic properties of cortical neurones
The callosal system of the superior parietal lobule in the monkey
No full textThe callosal connections of the superior parietal lobule, area 5 of Brodmann, were studied in macaque monkeys (M. nemestrina and M. fascicularis) using anatomical techniques based on both anterograde and retrograde axoplasmic transport of wheat-germ-agglutinin-conjugated horseradish peroxidase. From sagittal sections, two-dimensional flattened computer reconstructions of the volumes of cortical tissue containing callosal-projecting neurons (callosal efferent zone) and/or callosal terminal axons (callosal terminal territory) were obtained. Callosal zones were found in area 5, including the supplementaty sensory area, in a limited part of area 6, i.e., in the supplementary motor area, in area 7b, in the cortex of the dorsal bank of the sylvian fissure, and in a limited part of area 7a, in the cortex of the upper third of the rostral bank of the superior temporal sulcus. Callosal neurons in all cortical areas studied, though with regional variations, predominated in layer IIIb, but were also very numerous in layers VI and V. They were rare in other cortical laminae. In the cortical regions projecting heterotopically to area 5, the tangential distribution of callosal neurons was discontinuous because of the presence of large acallosal regions. These were not observed in area 5, although here the distribution of callosal neurons waxed and waned to the tangential cortical plane. Callosal axons to and/or from area 5 crossed the midline in the posterior, presplenial part of the corpus callosum. In the superior parietal lobule they terminated in radial patches or columns, spanning layers I-IV. These columns of various width (200-2,000 μm) were separated by gaps of similar size, free of such terminals. Callosal neurons were present not only within, but also between, the callosal terminal columns. Callosal neurons located within the callosal terminal columns were, in a statistically significant way, more numerous than those located between them. The callosal efferent zone occupied 71% of the tangential domain of area 5, whereas the callosal terminal territory occupied only 49% of it. This difference is statistically significant. The discontinuous columnar arrangement of callosal terminals and the periodic distribution of callosal neurons in the lateral part of area 5 defined three main bands of callosal connections of irregular shape which were oriented mediolaterally and ran parallel to the main architectonic borders, the border between areas 2 and 5 and that between 5 and 7
The anatomical substrate of callosal messages from SI and SII in the cat
No full textHorseradish peroxidase (HRP) was injected into the first (SI) or second (SII) somatosensory areas of 21 adult cats. The radial and tangential (normal and parallel to the pial surface, respectively) distribution and morphology of the callosal neurons were studied. HRP injections were combined with single unit recording in the contralateral cortex in order to determine which part of the somatosensory periphery is represented within the regions containing callosal neurons, the callosal (efferent) zones, in SI and SII. The callosal zone of SI extends over the trunk and part of the forepaw representation. In the forepaw and hindlimb representations callosal neurons projecting only to the contralateral SII are found, while in the trunk representation callosal neurons projecting to contralateral SI or SII are found. The callosal zone in SII extends widely throughout the forepaw representation in this area and projects to the contralateral SII but not to SI. In both SI and SII the collosal neurons are mainly located in layer III. A few of them are also found in layer VI. They are very rare in other layers. Callosal neurons in layer III are mostly pyramidal but exceptionally stellate; in layer VI they are pyramidal, triangular and occasionally stellate. These data indicate that transformations of the cortical somatosensory maps are achieved in the message sent through the corpus callosum. These transformations are i) determined by the extent and location of the callosal zones and perhaps by the distribution of callosal neurons within them, ii) different in different areas, iii) different in a same area, according to the cortical targets to which they are conveyed. The existence of callosal connections originated from areas of distal forepaw representation supplied a possible anatomical substrate for those types of intermanual transfer of tactile learning which depend upon the integrity of the corpus callosum
CORTICAL NETWORKS FOR VISUAL REACHING
No full textThe cortical anatomical substrates by which visual information may influence the frontal areas controlling reaching movements to visual targets were studied in monkeys. A reaching task was employed to characterize the arm-related regions of the frontal lobe. Injections of retrograde tracers into these physiologically defined cortical fields revealed a gradient of parallel cortico-cortical pathways originating in the superior parietal lobule and impinging upon different frontal regions. These results support the hypothesis that the superior parietal lobule can supply the frontal motor and premotor areas not only with the proprioceptive information but also with the visual input required for the control of reaching
The cortical network for eye–hand coordination and its relevance to understanding motor disorders of parietal patients
No full textCortical neurons in both superior (SPL) and inferior (IPL) parietal lobules are modulated by a variety of signals concerning planning and execution of eye and hand movement. Thanks to these properties, parietal neurons are ideally suited for eye–hand coordination during reaching. In SPL, a fundamental feature of neurons is the invariance of their directional tuning properties across tasks that require different forms of spatial relationships between the eye and the hand. In such conditions, the orientation of the preferred directions (PDs) of individual SPL cells cluster within a limited sector of space, the global tuning field (GTF), to be regarded as an ideal frame to dynamically match eye and hand signals on the basis of the orientation of their PDs. At the population level, the mean vectors of the GTF cover the direction continuum in a uniform fashion. These neurons are part of a parietal network richly interconnected with the premotor and motor areas of the frontal lobe. Thus, the reaching disorders of patients with optic ataxia might be interpreted as a consequence of the breakdown of the combinatorial mechanisms of the GTF of parietal neurons, and of their interplay with premotor cortex. In IPL, the main feature of eye and/or hand related neurons is the uneven distribution of their PDs, that mostly point toward the contralateral space. This anisotropy of the representation of directional motor space might explain the movement disorders that characterize directional hypokinesia in neglect patients. In conclusion, the study of the dynamic properties of parietal neurons and of their relationships with the premotor cortex via cortico-cortical connections provides a basis for an interpretation of movement disorders of parietal patients from a neurophysiological perspective
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