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Jon H Kaas

Publications and source records attributed to Jon H Kaas.

At least 19 recordsLinked to original sources

Myelin stains reveal an anatomical framework for the representation of the digits in somatosensory area 3b of macaque monkeys.

Brain sections cut parallel to the cortical surface revealed myelin-light septa that isolated representations of the digits and parts of the face, teeth, and tongue in area 3b of adult and infant macaque monkeys. The widths of the bands of cortex representing individual digits, as measured by the distances between isolating septa, were proportionally similar in infant (2-4 week) and adult monkeys. However, the bands for digits 1-3 were somewhat narrower in infant than adult monkeys. There was little variation in absolute widths across individuals in the infant or adult groups, or between left and right hemispheres of the same group. Widths for digits 1-4 progressively decreased. The results suggest that these isomorphs of digits emerge in prenatal or early postnatal development and typical variations in postnatal hand use have little impact on subsequent development. As the hand representation in somatosensory cortex of monkeys may be significantly altered after the partial loss of peripheral nerve inputs, the physiological representation is not completely constrained by the isolating septa. Instead, the septa may serve as a persistent marker of normal organization in studies of cortical reorganization.

Animals↗

Anatomical and functional organization of somatosensory areas of the lateral fissure of the New World titi monkey (Callicebus moloch).

The organization of anterior and lateral somatosensory cortex was investigated in titi monkeys (Callicebus moloch). Multiunit microelectrode recordings were used to identify multiple representations of the body, and anatomical tracer injections were used to reveal connections. (1) Representations of the face were identified in areas 3a, 3b, 1, S2, and the parietal ventral area (PV). In area 3b, the face was represented from chin/lower lip to upper lip and neck/upper face in a rostrocaudal sequence. The representation of the face in area 1 mirrored that of area 3b. Another face representation was located in area 3a. Adjoining face representations in S2 and PV exhibited mirror-image patterns to those of areas 3b and 1. (2) Two representations of the body, the rostral and caudal ventral somatosensory areas (VSr and VSc), were found in the dorsal part of the insula. VSc was roughly a reversal image of the S2 body representation, and VSr was roughly a reversal of PV. (3) Neurons in the insula next to VSr and VSc responded to auditory stimuli or to both auditory and somatosensory stimuli. (4) Injections of tracers within the hand representations in areas 3b, 1, and S2 revealed reciprocal connections between these three areas. Injections in areas 3b and 1 labeled the ventroposterior nucleus, whereas injections in S2 labeled the inferior ventroposterior nucleus. The present study demonstrates features of somatosensory cortex of other monkeys in titi monkeys, while revealing additional features that likely apply to other primates.

Animals↗

Bilateral effects of spinal overhemisections on the development of the somatosensory system in rats.

Connections of the forepaw regions of somatosensory cortex (S1) were determined in rats reared to maturity after spinal cord overhemisections at cervical level C3 on postnatal day 3. Overhemisections cut all ascending and descending pathways and intervening gray on one side of the spinal cord and the pathways of the dorsal funiculus contralaterally. Bilateral lesions of the dorsal columns reduced the size of the brainstem nuclei by 41%, and the ventroposterior lateral subnucleus (VPL) of the thalamus by 20%. Bilateral lesions also prevented the emergence of the normal cytochrome oxidase barrel pattern in forepaw and hindpaw regions of S1. Injections of wheat germ agglutinin conjugated to horseradish peroxidase were placed in the forepaw region of granular S1 and surrounding dysgranular S1 contralateral to the hemisection. The VPL nucleus was densely labeled, whereas the adjoining ventroposterior medial subnucleus, VPM, representing the head, was unlabeled. Thus, there was no evidence of abnormal connections of VPM to forepaw cortex. Foci of transported label in the ipsilateral hemisphere appeared to be in normal locations and of normal extents, but connections in the opposite hemisphere were broadly and nearly uniformly distributed in sensorimotor cortex in a pattern similar to that in postnatal rats. Rats with incomplete lesions that spared the dorsal column pathway on the left side but not the right demonstrated surprisingly normal distributions of callosal connections in the nondeprived right hemisphere, even though the injected left hemisphere was deprived. Thus, the development of the normal pattern of callosal connections depends on dorsal column input and not on normal interhemsipheric interactions.

Afferent Pathways↗

Optical imaging of visually evoked responses in prosimian primates reveals conserved features of the middle temporal visual area.

Optical imaging of intrinsic cortical responses to visual stimuli was used to characterize the organization of the middle temporal visual area (MT) of a prosimian primate, the bush baby (Otolemur garnetti). Stimulation with moving gratings revealed a patchwork of oval-like domains in MT. These orientation domains could, in turn, be subdivided into zones selective to directional movements that were mainly orthogonal to the preferred orientation. Similar, but not identical, zones were activated by movements of random dots in the preferred direction. Orientation domains shifted in preference systematically either around a center to form pinwheels or as gradual linear shifts. Stimuli presented in different portions of the visual field demonstrated a global representation of visual space in MT. As optical imaging has revealed similar features in MT of New World monkeys, MT appears to have retained these basic features of organization for at least the 60 million years since the divergence of prosimian and simian primates.

Animals↗

The visual pulvinar in tree shrews I. Multiple subdivisions revealed through acetylcholinesterase and Cat-301 chemoarchitecture.

Tree shrews are highly visual mammals closely related to primates. They have a large visual pulvinar complex, but its organization and relation to visual cortex is only partly known. We processed brain sections through the pulvinar with seven different procedures in an effort to reveal histologically distinct compartments. The results revealed three major subdivisions. A dorsal subdivision, Pd, stains darkly for acetylcholinesterase (AChE) and occupies the dorsoposterior one-third of the pulvinar complex. A ventral subdivision, Pv, stains darkly when processed with the Cat-301 antibody and occupies the ventroanterior fifth of the pulvinar complex along the brachium of the superior colliculus. Unexpectedly, part of Pv is ventral to the brachium. A large central subdivision, Pc, stains moderately dark for AChE and cytochrome oxidase (CO), and very light for Cat-301. Pc includes about half of the pulvinar complex, with parts on both sides of the brachium of the superior colliculus. These architectonic results demonstrate that the pulvinar complex of tree shrews is larger and has more subdivisions than previously described. The complex resembles the pulvinar of primates by having a portion ventral to the brachium and by having histochemically distinct nuclei; the number of nuclei is less than in primates, however.

Acetylcholinesterase↗

The visual pulvinar in tree shrews II. Projections of four nuclei to areas of visual cortex.

Patterns of thalamocortical connections were related to architectonically defined subdivisions of the pulvinar complex and the dorsolateral geniculate nucleus (LGN) in tree shrews (Tupaia belangeri). Tree shrews are of special interest because they are considered close relatives of primates, and they have a highly developed visual system. Several distinguishable tracers were injected within and across cortical visual areas in individual tree shrews in order to reveal retinotopic patterns and cortical targets of subdivisions of the pulvinar. The results indicate that each of the three architectonic regions of the pulvinar has a distinctive pattern of cortical connections and that one of these divisions is further divided into two regions with different patterns of connections. Two of the pulvinar nuclei have similar retinotopic patterns of projections to caudal visual cortex. The large central nucleus of the pulvinar (Pc) projects to the first and second visual areas, V1 and V2, and an adjoining temporal dorsal area (TD) in retinotopic patterns indicating that the upper visual quadrant is represented dorsal to the lower quadrant in Pc. The smaller ventral nucleus (Pv) which stains darkly for the Cat-301 antigen, projects to these same cortical areas, with a retinotopic pattern. Pv also projects to a temporal anterior area, TA. The dorsal nucleus (Pd), which densely expresses AChE, projects to posterior and ventral areas of temporal extrastriate cortex, areas TP and TPI. A posterior nucleus, Pp, projects to anterior areas TAL and TI, of the temporal lobe, as well as TPI. Injections in different cortical areas as much as 6 mm apart labeled overlapping zones in Pp and double-labeled some cells. These results indicate that the visual pulvinar of tree shrews contains at least four functionally distinct subdivisions, or nuclei. In addition, the cortical injections revealed that the LGN projects topographically and densely to V1 and that a significant number of LGN neurons project to V2 and TD.

Acetylcholinesterase↗

Patterned activity via spinal dorsal quadrant inputs is necessary for the formation of organized somatosensory maps.

The normal development of the somatosensory system requires intact sensory inputs from the periphery during a critical window of time early in development. Here we determined how the removal of only part of the ascending spinal inputs early in development affects the anatomical and neurophysiological development of the somatosensory system. We performed spinal overhemisections in rat pups at C3/C4 levels on the third day after birth. This procedure hemisects the spinal cord on one side and transects the dorsal funiculus on the other side. When the rats were 6-8 months old, the responsiveness and somatotopy of the primary somatosensory cortex (S1) contralateral to the hemisection were determined using standard multiunit mapping techniques. Sections of the flattened cortex were processed for cytochrome oxidase activity, Nissl substance, or myelin. We found that histologically apparent modules that are normally present in the regions of the forepaw and the hindpaw representations were absent, whereas the lateral barrel field representing the face was completely normal. The neurons in the forepaw regions of S1 either did not respond to the stimulation of the skin of any region of the body or responded to the stimulation of the upper arm afferents that enter the spinal cord rostral to the site of the lesion. The results show that a lack of normal sensory inputs via ascending pathways in the dorsal spinal cord during early development results in massive anatomical and neurophysiological abnormalities in the cortex. Intact crossed spinothalamic pathways are unable to support the normal development of the forepaw barrels.

Afferent Pathways↗

Responses of neurons in the middle temporal visual area after long-standing lesions of the primary visual cortex in adult new world monkeys.

The retinotopic organization of the middle temporal visual area (MT) was determined in six adult owl monkeys and one adult marmoset 69 d to 10 months after lesions of the dorsolateral primary visual cortex (V1). The lesions removed were limited to extensive parts of the representation of the lower visual quadrant in V1. Microelectrodes were used to record from neurons at numerous sites in MT to determine whether parts of MT normally devoted to the lower visual quadrant (1) were unresponsive to visual stimuli, (2) acquired responsiveness to inputs from intact portions of V1, or (3) became responsive to some other visually driven input such as a relay from the superior colliculus via the pulvinar to MT. All monkeys (n = 6) with moderate to moderately large lesions had unresponsive portions of MT even after 10 months of recovery. These unresponsive regions were retinotopically equivalent to the removed parts of V1 in normal animals. Thus, there was no evidence for an alternative source of activation. In addition, these results indicate that any retinotopic reorganization of MT based on inputs from intact portions of V1 was not extensive, yet neurons near the margins of responsive cortex may have acquired new receptive fields, and the smallest 5 degrees lesion of V1 failed to produce an unresponsive zone. Deprived portions of MT were not remarkably changed in histological appearance in cytochrome oxidase, Nissl, and Wisteria floribunda agglutinin preparations. Nevertheless, some reduction in myelin staining and other histological changes were suggested. We conclude that MT is highly dependent on V1 for activation in these monkeys, and alternative sources do not become effective over months when normal activation is absent. Additionally, remaining V1 inputs have only a limited capacity to expand their activation territory into deprived portions of MT.

Animals↗

Somatosensory cortex of prosimian Galagos: physiological recording, cytoarchitecture, and corticocortical connections of anterior parietal cortex and cortex of the lateral sulcus.

Compared with our growing understanding of the organization of somatosensory cortex in monkeys, little is known about prosimian primates, a major branch of primate evolution that diverged from anthropoid primates some 60 million years ago. Here we describe extensive results obtained from an African prosimian, Galago garnetti. Microelectrodes were used to record from large numbers of cortical sites in order to reveal regions of responsiveness to cutaneous stimuli and patterns of somatotopic organization. Injections of one to several distinguishable tracers were placed at physiologically identified sites in four different cortical areas to label corticortical connections. Both types of results were related to cortical architecture. Three systematic representations of cutaneous receptors were revealed by the microelectrode recordings, S1 proper or area 3b, S2, and the parietal ventral area (PV), as described in monkeys. Strips of cortex rostral (presumptive area 3a) and caudal (presumptive area 1-2) to area 3b responded poorly to tactile stimuli in anesthetized galagos, but connection patterns with area 3b indicated that parallel somatosensory representations exist in both of these regions. Area 3b also interconnected somatotopically with areas S2 and PV. Areas S2 and PV had connections with areas 3a, 3b, 1-2, each other, other regions of the lateral sulcus, motor cortex (M1), cingulate cortex, frontal cortex, orbital cortex, and inferior parietal cortex. Connection patterns and recordings provided evidence for several additional fields in the lateral sulcus, including a retroinsular area (Ri), a parietal rostral area (PR), and a ventral somatosensory area (VS). Galagos appear to have retained an ancestral preprimate arrangement of five basic areas (S1 proper, 3a, 1-2, S2, and PV). Some of the additional areas suggested for lateral parietal cortex may be primate specializations.

Animals↗

Somatosensory input to the ventrolateral thalamic region in the macaque monkey: potential substrate for parkinsonian tremor.

In the present study, we determined the anatomic relationships between somatosensory and motor pathways within ventrolateral (VL) thalamic nuclei of the motor thalamus of macaque monkeys. In labeling experiments, four macaque monkeys (Macaca mulatta) received injections of biotinylated dextran amine and wheat germ agglutinin conjugated to horseradish peroxidase into the cerebellar nuclei or internal segment of the globus pallidus and cervical segments of the spinal cord, respectively. Each tracer was visualized in brain sections by sequentially using a different chromogen. Labeled terminals were plotted and superimposed on adjacent brain sections processed for Nissl substance, acetylcholinesterase, and the antigens for calbindin and Cat-301 to reveal thalamic nuclei. The labeled cerebellar terminals were distributed throughout the posterior VL (VLp), whereas the labeled pallidothalamic terminals were concentrated in the anterior VL and the ventral anterior nucleus. The spinothalamic input was directed mostly to the ventral posterior complex and cells just caudal to it. In addition, the patches of spinothalamic terminations intermingled and partly overlapped with the cerebellothalamic, but not with the pallidothalamic terminations within VLp. The regions of overlap of somatosensory and cerebellar inputs within the VLp of the present study appear to correspond to the reported locations of the tremor-related cells in parkinsonian patients. Thus, the overlapping spinothalamic and cerebellar inputs may provide a substrate for the altered activity of motor thalamic neurons in such patients.

Acetylcholinesterase↗

Few intrinsic connections cross the hand-face border of area 3b of New World monkeys.

After long-standing loss of afferents from the hand, the hand representation in area 3b of the somatosensory cortex of monkeys becomes responsive to touch on the face. Because the reactivation of deprived hand cortex by the face inputs could depend on axonal connections across the hand-face border, we determined the extent of such connections in New World marmosets, owl monkeys, and squirrel monkeys. Small injections of anatomic tracers were placed in the hand or the face representations after these representations were identified by microelectrode recordings. The positions of retrogradely labeled neurons were plotted in processed brain sections cut parallel to the brain surface, and their locations were related to anatomic isomorphs of the hand and face representations revealed in adjacent brain sections stained for myelin. In these sections, the hand-face border was clearly visualized as a myelin-poor septum. The intrinsic connections of area 3b labeled by injections in either the hand or face representations were almost completely confined to their respective representation, and very few neurons projected across the border. In addition, neurons in the somatosensory thalamus labeled by injections in either face or hand representations were confined to either VPM, representing the face, or the hand subnucleus of VPL. Thus the reactivation of hand cortex by face stimulation does not depend on a previously existing network of intrinsic cortical connections across the hand-face border, or mismatched thalamocortical projections.

Afferent Pathways↗

Optical imaging reveals retinotopic organization of dorsal V3 in New World owl monkeys.

Optical imaging of intrinsic responses to visual stimuli in extrastriate cortex of owl monkeys provided evidence for the dorsal half of the third visual area, V3. Visual stimuli were used to selectively activate locations in dorsolateral V2 and the rostrally adjoining presumptive V3. Consistent with the proposed retinotopies of dorsal V2 and dorsal V3, small bars of drifting gratings along the horizontal meridian of the contralateral hemifield activated cortex along the V2V3 border, whereas such stimuli along the vertical meridian activated cortex along the rostral border of V3. Stimuli in limited locations in the lower visual quadrant revealed mirror reversals of retinotopy in dorsal V2 and V3, whereas stimuli in the upper visual quadrant failed to activate either region. Brain sections processed for cytochrome oxidase from the same cases provided architectonic borders of V2 that matched those indicated by the optical imaging. The results support the concept that a narrow dorsal V3 exists in monkeys. V3d borders dorsal V2 and contains a smaller, mirror-image representation of the lower visual quadrant.

Action Potentials↗

Pulvinar and other subcortical connections of dorsolateral visual cortex in monkeys.

The present study used injections of neuroanatomical tracers to determine the subcortical connections of the caudal and rostral subdivisions of the dorsolateral area (DL) and the middle temporal crescent area (MT(C)) in owl monkeys (Aotus trivirgatus), squirrel monkeys (Saimiri sciureus), and macaque monkeys (Macaca fascicularis and M. radiata). Emphasis was on connections with the pulvinar. Patterns of corticopulvinar connections were related to subdivisions of the inferior pulvinar (PI) defined by histochemical or immunocytochemical architecture. Connections of DL/MT(C) were with the PI subdivisions, PICM, PICL, and PIp; the lateral pulvinar (PL); and, more sparsely, the lateral portion of the medial pulvinar (PM). In squirrel monkeys, there was a tendency for caudal DL to have stronger connections with PICL than PICM and for rostral DL/MT(C) to have stronger connections with PICM than PICL. In all three primates, DL/MT(C) had reciprocal connections with the pulvinar and claustrum; received afferents from the locus coeruleus, dorsal raphe, nucleus annularis, central superior nucleus, pontine reticular formation, lateral geniculate nucleus, paracentral nucleus, central medial nucleus, lateral hypothalamus, basal nucleus of the amygdala, and basal nucleus of Meynert/substantia innominata; and sent efferents to the pons, superior colliculus, reticular nucleus, caudate, and putamen. Projections from DL/MT(C) to the nucleus of the optic tract were also observed in squirrel and owl monkeys. Similarities in the subcortical connections of the dorsolateral region, especially those with the pulvinar, provide further support for the conclusion that the DL regions are homologous in the three primate groups.

Animals↗

Evidence from V1 connections for both dorsal and ventral subdivisions of V3 in three species of New World monkeys.

We used patterns of connections of primary visual cortex (V1) to reevaluate differing proposals on the organization of extrastriate cortex in three species of New World monkeys. Several fluorescent tracers and the bidirectional tracer cholera toxin B subunit (CTB) were injected into dorsal V1 (representing the lower visual quadrant) and ventral V1 (representing the upper visual quadrant) of titi, squirrel, and owl monkeys. Labeled cells and terminals were plotted on brain sections cut parallel to the surface of flattened cortex and were related to architectonic boundaries. The results provided compelling evidence for both dorsal V3 with dorsal V1 connections and ventral V3 with ventral V1 connections. The connection pattern indicated that V3 represents the visual hemifield as a mirror image of V2. In addition, V3 could be recognized by a weak banding pattern in brain sections processed for cytochrome oxidase. V1 has connections with at least 12 subdivisions of visual cortex, with half of the connections involving V2 and 20% V3. Comparable results were obtained from all three species, suggesting that visual cortex is similarly organized.

Animals↗

Cortical and thalamic connections of the parietal ventral somatosensory area in marmoset monkeys (Callithrix jacchus).

Microelectrode mapping methods were used to define the parietal ventral somatosensory area (PV) on the upper bank of the lateral sulcus in five marmosets (Callithrix jacchus). In the same animals, neuroanatomical tracers were placed into electrophysiologically identified sites in PV and/or the second somatosensory area (S2). Foci of anterograde and retrograde label were related to electrophysiological maps of cortical areas and cortical and thalamic architecture. The results lead to the following conclusions: (1) Multiunit recordings from cortex on the upper bank of the lateral sulcus demonstrate that PV is somatotopically organized, with the face representation adjoining area 3b and the hindlimb and tail representations away from this border in cortex deep on the upper bank of the lateral sulcus. The forelimb representation is caudal in PV adjacent to the S2 forelimb representation. The body surface representation in PV approximates a mirror image of that in S2; (2) Areas PV and S2 are less myelinated and have less cytochrome oxidase enzyme activity than area 3b; (3) The ventroposterior inferior nucleus (VPI) of the thalamus provides the major somatosensory projections to PV. PV is reciprocally connected with VPI and anterior pulvinar; (4) PV has ipsilateral cortical connections with areas 3a, 3b, 1, and M1 and higher order somatosensory fields, and at least most of these connections are somatotopically matched; and (5) Callosal connections of PV are with S2 and PV of the other cerebral hemisphere. These results further establish PV as one of at least four somatosensory areas of the lateral sulcus of primates.

Animals↗