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Prefrontal connections of the parabelt auditory cortex in macaque monkeys.

In the present study, we determined connections of three newly defined regions of auditory cortex with regions of the frontal lobe, and how two of these regions in the frontal lobe interconnect and connect to other portions of frontal cortex and the temporal lobe in macaque monkeys. We conceptualize auditory cortex as including a core of primary areas, a surrounding belt of auditory areas, a lateral parabelt of two divisions, and adjoining regions of temporal cortex with parabelt connections. Injections of several different fluorescent tracers and wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) were placed in caudal (CPB) and rostral (RPB) divisions of the parabelt, and in cortex of the superior temporal gyrus rostral to the parabelt with parabelt connections (STGr). Injections were also placed in two regions of the frontal lobe that were labeled by a parabelt injection in the same case. The results lead to several major conclusions. First, CPB injections label many neurons in dorsal prearcuate cortex in the region of the frontal eye field and neurons in dorsal prefrontal cortex of the principal sulcus, but few or no neurons in orbitofrontal cortex. Fine-grain label in these same regions as a result of a WGA-HRP injection suggests that the connections are reciprocal. Second, RPB injections label overlapping prearcuate and principal sulcus locations, as well as more rostral cortex of the principal sulcus, and several locations in orbitofrontal cortex. Third, STGr injections label locations in orbitofrontal cortex, some of which overlap those of RPB injections, but not prearcuate or principal sulcus locations. Fourth, injections in prearcuate and principal sulcus locations labeled by a CPB injection labeled neurons in CPB and RPB, with little involvement of the auditory belt and no involvement of the core. In addition, the results indicated that the two frontal lobe regions are densely interconnected. They also connect with largely separate regions of the frontal pole and more medial premotor and dorsal prefrontal cortex, but not with the extensive orbitofrontal region which has RPB and STGr connections. The results suggest that both RPB and CPB provide the major auditory connections with the region related to directing eye movements towards stimuli of interest, and the dorsal prefrontal cortex for working memory. Other auditory connections to these regions of the frontal lobe appear to be minor. RPB has connections with orbitofrontal cortex, important in psychosocial and emotional functions, while STGr primarily connects with orbital and polar prefrontal cortex.

Animals↗

Topographic organization in the corticocortical connections of medial agranular cortex in rats.

Medial agranular cortex (AGm) is a narrow, longitudinally oriented region known to have extensive corticortical connections. The rostral and caudal portions of AGm exhibit functional differences that may involve these connections. Therefore we have examined the rostrocaudal organization of the afferent cortical connections of AGm by using fluorescent tracers, to determine whether there are significant differences between rostral and caudal AGm. Mediolateral patterns have also been examined in order to compare the pattern of corticocortical connections of AGm to those of the laterally adjacent lateral agranular cortex (AGl) and medially adjacent anterior cingulate area (AC). In the rostrocaudal domain, there are notable patterns in the connections of AGm with somatic sensorimotor, visual, and retrosplenial cortex. Rostral AGm receives extensive afferents from the caudal part of somatic sensorimotor area Par I, whereas caudal AGm receives input largely from the hindlimb cortex (area HL). Middle portions of AGm show an intermediate condition, indicating a continuously changing pattern rather than the presence of sharp border zones. The whole of the second somatic sensorimotor area Par II projects to rostral AGm, whereas caudal AGm receives input only from the caudal portion of Par II. Visual cortex projections to AGm originate in areas Oc1, Oc2L and Oc2M. Connections of rostral AGm with visual cortex are noticeably less dense than those of mid and caudal AGm, and are focused in area Oc2L. The granular visual area Oc1 projects almost exclusively to mid and caudal AGm. Retrosplenial cortex has more extensive connections with caudal AGm than with rostral AGm, and the agranular and granular retrosplenial subregions are both involved. Other cortical connections of AGm show little or no apparent rostrocaudal topography. These include afferents from orbital, perirhinal, and entorhinal cortex, all of which are bilateral in origin. In the mediolateral dimension, AGm has more extensive corticocortical connections than either AGl or AC. Of these three neighboring areas, only AGm has connections with the somatic sensorimotor, visual, retrosplenial and orbital cortices. In keeping with its role as primary motor cortex, AGl is predominantly connected with area Par I of somatic sensorimotor cortex, specifically rostral Par I. AGl receives no input from visual or retrosplenial cortex. Anterior cingulate cortex has connections with visual area Oc2 and with retrosplenial cortex, but none with somatic sensorimotor cortex. Orbital cortex projections are sparse to AGl and do not appear to involve AC.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Susceptibility of different cell layers of the anterior and posterior part of the piriform cortex to electrical stimulation and kindling: comparison with the basolateral amygdala and "area tempestas".

Several lines of evidence suggest that the piriform cortex functions as a generator in the development and propagation of forebrain (limbic type) seizures, particularly in the kindling model of epilepsy. It is, however, not clear where, within the rather large piriform cortex region, the generator resides, and how much tissue is involved. Highly sensitive loci to chemical or electrical stimulation have been described both in the deep anterior and posterior parts of the piriform cortex. Furthermore, data from piriform cortex slice preparations indicated that epileptiform potentials originate in deep structures, particularly the endopiriform nucleus that underlies the piriform cortex. In the present study, in rats, we implanted stimulation and recording electrodes in various rostrocaudal locations of the piriform cortex and endopiriform nucleus, including the "area tempestas", i.e. a structure in the anterior part of the piriform cortex previously proposed to be critically involved in the generation of convulsive seizures of limbic origin. Within the piriform cortex, electrodes were aimed at different cellular layers of this structure. For comparison, additional animals received electrodes in different parts of the basolateral amygdala. A total of 19 different locations was obtained in this way. The susceptibility of these locations to electrical stimulation was characterized by determining the threshold for induction of afterdischarges. The afterdischarge threshold was lowest in layer III of the posterior piriform cortex and some locations in the endopiriform nucleus, whereas amygdala and "area tempestas" displayed higher values. In several animals, particularly those with electrodes in layer III of the posterior piriform cortex, spontaneous spiking was seen in prestimulation recordings, whereas this was never observed in recordings from the amygdala. Subsequent kindling by repeated stimulation of the various locations demonstrated marked differences in afterdischarge threshold reduction and kindling rate. The most marked decreases in afterdischarge threshold were seen in locations within layer III of the piriform cortex, whereas several other locations, including the "area tempestas", exhibited only moderate decreases or no decrease at all. In contrast to previous observations with only few locations in the piriform cortex region, the posterior piriform cortex was not in general slower to kindle than the anterior piriform cortex, although some locations in the posterior piriform cortex exhibited significantly lower kindling rates than the amygdala. The highest kindling rate was seen in the dorsal endopiriform nucleus.(ABSTRACT TRUNCATED AT 400 WORDS)

Amygdala↗

Cortical somatosensory evoked potentials. II. Effects of excision of somatosensory or motor cortex in humans and monkeys.

1. To clarify the generators of human short-latency somatosensory evoked potentials (SEPs) thought to arise in sensorimotor cortex, we studied the effects on SEPs of surgical excision of somatosensory or motor cortex in humans and monkeys. 2. Normal median nerve SEPs (P20-N30, N20-P30, and P25-N35) were recorded from the cortical surface of a patient (G13) undergoing a cortical excision for relief of focal seizures. All SEPs were abolished both acutely and chronically after excision of the hand area of somatosensory cortex. Similarly, excision of the hand area of somatosensory cortex abolished corresponding SEPs (P10-N20, N10-P20, and P12-N25) in monkeys. Excision of the crown of monkey somatosensory cortex abolished P12-N25 while leaving P10-N20 and N10-P20 relatively unaffected. 3. After excision of the hand area of motor cortex, all SEPs were present when recorded from the cortical surface of a patient (W1) undergoing a cortical excision for relief of focal seizures. Similarly, all SEPs were present in monkeys after excision of the hand area of motor cortex. 4. Although all SEPs were present after excision of motor cortex in monkeys, variable changes were observed in SEPs after the excisions. However, these changes were not larger than the changes observed after excision of parietal cortex posterior to somatosensory cortex. We concluded that the changes were not specific to motor cortex excision. 5. These results support two major conclusions. 1) Median nerve SEPs recorded from sensorimotor cortex are produced by generators in two adjacent regions of somatosensory cortex: a tangentially oriented generator in area 3b, which produces P20-N30 (human) and P10-N20 (monkey) [recorded anterior to the central sulcus (CS)] and N20-P30 (human) and N10-P20 (monkey) posterior to the CS; and a radially oriented generator in area 1, which produces P25-N35 (human) and P12-N25 (monkey) recorded from the postcentral gyrus near the CS. 2) Motor cortex makes little or no contribution to these potentials.

Adolescent↗

The topography of the mesencephalic and pontine projections from the cingulate cortex of the rat.

A projection from the rat midline cortex to the midbrain and pons has been recognized for several years. The present study is a detailed analysis of this projection using the autoradiographic technique. Small injections of [3H]amino acids were placed within individual segments of the cingulate cortex in 68 rats. The resulting material reaffirmed the existence of the cingulo-brainstem projections and demonstrated that a precise topographical relationship exists between the cingulate cortex cells of origin and their termination fields within the brainstem. The most ventral and anterior segment of the cortex (IRaa) projects to the ventral periaqueductal gray, to the dorso-medial ventral pontine nuclei and to the lateral tegmental region. Conversely, the dorso-anterior cortex (IRca) projects to the superior colliculus, the dorso-lateral periaqueductal gray, and the medio-ventral ventral pontine nuclei. The intermediate anterior cortex projects to both dorsal and ventral periaqueductal gray, lightly to the superior colliculus, and to the medio-intermediate ventral pontine nuclei. The posterior half of the infraradiata (IR) cortex projects to the dorso-lateral periaqueductal gray, to the superior colliculus, and to the region of the ventral pontine nuclei slightly lateral to the terminal zone occupied by the anterior IR cortex. Increasingly dorsal segments of the IR beta cortex project to more increasingly ventral areas of the ventral nuclei. The posterior portion of the midline cortex (retrosplenial cortex, R) does not project to the dorsal midbrain, but it does topographically project to the ventral pontine nuclei, lateral to the terminal zone of the IR axons. Increasingly, posterior regions of the R cortex project to more lateral regions of the ventral nuclei, and increasingly, dorsal cells of the R cortex project to more dorsal regions of the ventral nuclei. These data demonstrate a very precise topography of brainstem projections which may underlie the visceral and somatic motor functions of the cingulate cortex, as well as the ability of the cingulate cortex to modulate sensory information and emotional behavior.

Animals↗

Associative synaptic potentials in the piriform cortex of the isolated guinea-pig brain in vitro.

The involvement of local and remote associative fibres in the generation of piriform cortex synaptic potentials was investigated in the isolated guinea-pig brain maintained in vitro by arterial perfusion by implementing current source density analysis (CSD) on cortical field potential profiles. Previous hypotheses were verified using acute surgical isolation of piriform cortical areas to study different synaptic events separately. Stimulation of the lateral olfactory tract activated associative potentials throughout the piriform cortex. In the anterior piriform cortex, the current sinks responsible for the generation of associative potentials were located in the superficial portion of layer Ib and in layer III. In the posterior piriform cortex, two associative events were observed: an early sink located in the superficial part of layer Ib, followed by a sink in the deep part of the same layer. In the anterior piriform cortex, local associative synaptic potentials were separated from the component carried by long projective fibres by surgically isolating a small area of cortex monosynaptically activated by lateral olfactory tract stimulation. In this patch of lateral olfactory tract-connected anterior piriform cortex, local associative sinks were observed in the superficial Ib layer and in layer III. Monosynaptic activation of the isolated patch of anterior piriform cortex induced purely associative potentials throughout the piriform cortex. These potentials were mediated by the synaptic activation of apical dendrites in the superficial Ib layer and selectively abolished by severing the long associative fibres. The anterior piriform cortex layer III sink and the posterior piriform cortex deep Ib associative component were evoked by the activation of large population spikes in the monosynaptic anterior piriform cortex and the disynaptic posterior piriform cortex response respectively. These two sinks are presumably generated locally through a polysynaptic circuit, whose activation depends on the degree of cortical excitation. Olfactory signal processing in the guinea-pig piriform cortex during states of normal excitability is supported by the interactions between associative inputs impinging on the synapses located separately on the dendrites of pyramidal neurons. An increase in the synchronization of piriform cortex neuron discharge activates usually silent local circuit synapses.

Afferent Pathways↗

Contribution of thalamic input to the specification of cytoarchitectonic cortical fields in the primate: effects of bilateral enucleation in the fetal monkey on the boundaries, dimensions, and gyrification of striate and extrastriate cortex.

Bilateral enucleation was performed at different fetal ages during corticogenesis, and the brains were prepared for histological examination. Early-enucleated fetuses (operated prior to embryonic day 77) showed morphological changes at the level of the thalamus and the cortex. In the thalamus, there was a loss of lamination and a decrease in size of the lateral geniculate nucleus. There was a decrease in the size of the inferior pulvinar, but there was no change in the lateral pulvinar. The border of striate cortex was as sharp in the enucleates as it was in the normal monkeys. In three of the four early enucleates, we observed an interdigitation of striate and extrastriate cortex. In three of the early enucleates, we observed a small island of nonstriate cortex near the striate border that was surrounded entirely by striate cortex. Enucleation led to an age-related reduction of striate cortex. This reduction was greater in the operculum than in the calcarine fissure. The reduction of striate cortex was accompanied by an increase in the dimensions of extrastriate visual cortex, so that the overall dimensions of the neocortex remained invariant. The extrastriate cortex in the enucleated animals presented a uniform cytoarchitecture and was indistinguishable from area 18 in the normal animal. There were changes in the gyral pattern that were restricted mainly to the cortex on the operculum. A deepening of minor dimples as well as the induction of a variable number of supplementary sulci led to an increase in the convolution of the occipital lobe. These results are discussed with respect to the specification of cortical areas. They demonstrate that the reduction in striate cortex was not accompanied by an equivalent reduction in the neocortex; rather, there was a border shift, and a large volume of cortex that was destined to become striate cortex appears to be cytoarchitectonically normal extrastriate cortex.

Aging↗

Macaque monkey retrosplenial cortex: II. Cortical afferents.

We investigated the cortical afferents of the retrosplenial cortex and the adjacent posterior cingulate cortex (area 23) in the macaque monkey by using the retrograde tracers Fast blue and Diamidino yellow. We quantitatively analyzed the distribution of labeled neurons throughout the cortical mantle. Injections involving the retrosplenial cortex resulted in labeled neurons within the retrosplenial cortex and in areas 23 and 31 (approximately 78% of the total labeled cells). In the remainder of the cortex, the heaviest projections originated in the hippocampal formation, including the entorhinal cortex, subiculum, presubiculum, and parasubiculum. The parahippocampal and perirhinal cortices also contained many labeled neurons, as did the prefrontal cortex, mainly in areas 46, 9, 10, and 11, and the occipital cortex, mainly area V2. Injections in area 23 also resulted in numerous labeled cells in the posterior cingulate and retrosplenial regions (approximately 67% of total labeled cells). As in the retrosplenial cortex, injections of area 23 led to many labeled neurons in the frontal cortex, although most of these cells were in areas 9 and 46. Larger numbers of retrogradely labeled cells were also distributed more widely in the posterior parietal cortex, including areas 7a, 7m, LIP, and DP. There were some labeled cells in the parahippocampal cortex. These connections are consistent with the retrosplenial cortex acting as an interface between the working memory functions in the prefrontal areas and the long-term memory encoding in the medial temporal lobe. The posterior cingulate cortex, in contrast, may be more highly associated with visuospatial functions.

Afferent Pathways↗

Connections of the visual cortex in the hedgehog (Paraechinus hypomelas). II. Corticocortical projections.

Cortical subdivisions based on cytoarchitectural and myeloarchitectural observations of normal tissue were correlated with the topography of corticocortical connections in the visual system of the Pakistani hedgehog. Large subpial aspiration lesions were made in both visual and non-visual cortical regions to determine the areal limits of the corticocortical connections of the visual cortex. Subsequently, discrete electrolytic lesions were placed within the visual cortex. After appropriate survival periods, the brains were processed and stained with the Fink-Heimer technique. The results of these studies show that the visual cortex may be subdivided into four distinct regions from lateral to medial; the lateral parastriate cortex, the lateral and medial part of striate cortex, and the medial parastriate cortex. Within these regions, interhemispheric connections between visual cortices arise mainly in the lateral striate and lateral parastriate regions and terminate in a single band within the lateral portion of the cytoarchitecturally defined striate cortex. These corticocortical projections, therefore, substantially overlap with the geniculostriate projections. Lateral striate cortex and lateral parastriate cortex project in a reciprocal fashion that correlates well with the physiologically defined mirror image representation of two retinotopic maps of the binocular visual field on cortex. These connections are reflected about a line that is closely correlated with the medial edge of the band of commissural axon terminals that is located within the lateral striate cortex, instead of corresponding exactly with the striate-parastriate border as they do in other mammals. Medial striate cortex projects to medial parastriate cortex, indicating that the monocular portion of V I is related to a separate secondary area of cortex on the medial wall of the hemisphere.

Animals↗

Regional distribution of cholecystokinin receptors in primate cerebral cortex determined by in vitro receptor autoradiography.

Cholecystokinin (CCK) is a putative peptide neurotransmitter present in high concentration in the cerebral cortex. By using techniques of in vitro receptor autoradiography, CCK binding sites in primate cortex were labeled with 125I-Bolton-Hunter-labeled CCK-33 (the 33-amino-acid C-terminal peptide) and 3H-CCK-8 (the C-terminal octapeptide). Biochemical studies performed on homogenized and slide-mounted tissue sections showed that the two ligands labeled a high-affinity, apparently single, saturable site. Autoradiography revealed that binding sites labeled by both ligands were anatomically indistinguishable and were distributed in two basic patterns. A faint and diffuse label characterized portions of medial prefrontal cortex, premotor and motor cortices, the superior parietal lobule, and the temporal pole. In other cortical areas the pattern of binding was layer-specific; i.e., binding sites were concentrated within particular cortical layers and were superimposed upon the background of diffuse label. Layer-specific label was found in the prefrontal cortex, anterior and posterior cingulate gyrus, somatosensory cortex, inferior parietal lobule, retrosplenial cortex, insula, temporal lobe cortices, and in the primary visual and adjacent visual association cortices. The areal and laminar localization of layer-specific CCK binding sites consistently coincided with the cortical projections of thalamic nuclei. In prefrontal cortex, CCK binding sites were present in layers III and IV, precisely paralleling the terminal fields of thalamocortical projections from the mediodorsal and medial pulvinar nucleus of the thalamus. In somatosensory cortex, the pattern of CCK binding in layer IV coincided with thalamic inputs arising from the ventrobasal complex, while in the posterior cingulate gyrus, insular cortex, and retrosplenial cortex, layer IV and lower III binding mirrored the laminar distribution of cortical afferents of the medial pulvinar. CCK binding in layers IVa, IVc alpha, IVc beta, and VI of primary visual cortex corresponded to the terminal field disposition of lateral geniculate neurons, whereas in adjacent visual association cortex, binding in layers III, IV, and VI faithfully followed the cortical distribution of projections from the inferior and lateral divisions of the pulvinar nucleus of the thalamus. We interpret the diffusely labeled binding sites in primate cortex as being associated with the intrinsic system of CCK-containing interneurons that are distributed throughout all layers and areas of the cortex. The stratified binding sites, however, appear to be associated with specific extrinsic peptidergic projections.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Number and size of neurons and synapses in the motor cortex of cats raised in different environmental complexities.

In a previous study we have shown that the richness of the environment affects the number of neurons, the size of their nuclei, the number of round-asymmetrical synapses per neuron, the numerical density (number per unit volume; NV) of flat-symmetrical synaptic contacts, their number per neuron and their size in the visual cortex of cats. Of these, the number of flat-symmetrical synapses per unit volume is particularly affected (there are nearly twice as many per mm3 in the impoverished cortex). Several studies in the rat have shown that environmentally induced changes in cortical thickness occur in the occipital regions but are much smaller or absent in the frontal regions. In order to determine if the cat motor cortex is also resistant to environmental changes, we have estimated the number and size of neurons and of synapses in individual laminae of motor cortex, area 4 gamma, in six pairs of cats raised either in a colony (EC: enriched condition) or in isolation (IC: impoverished condition). For the neurons, we have found that the numerical density (28,900 neurons per mm3 of EC and 29,500 neurons/mm3 of IC motor cortex), the number under 1 mm2 of cortical surface (49,400 and 49,200 in EC and IC cats), and the size of the neuronal nuclei (82 vs 80 microns2 in EC and IC animals) were not significantly affected. The number of flat-symmetrical synapses per neuron (1,470 in EC vs 1,400 in IC cortex), their size (0.33 micron in both groups) and even their number per unit volume, which was so greatly affected in the visual cortex, remains unchanged (43 million/mm3 and 41 million/mm3 in EC and IC motor cortex). We did find however, a significant difference (p less than 0.05) in the numerical density of round-asymmetrical synapses which is 13% greater in the impoverished motor cortex (216 million/mm3 in EC vs 247 million/mm3 in IC cortex). Our results confirm that the motor cortex is much less affected by the richness of the environment than the visual cortex: In fact, the cat motor cortex is hardly affected at all. Furthermore our results represent the most complete data presently available on the number and size of neurons and synapses in individual laminae of the cat motor cortex.

Animals↗

Connections between the retrosplenial cortex and the hippocampal formation in the rat: a review.

The retrosplenial cortex is situated at the crossroads between the hippocampal formation and many areas of the neocortex, but few studies have examined the connections between the hippocampal formation and the retrosplenial cortex in detail. Each subdivision of the retrosplenial cortex projects to a discrete terminal field in the hippocampal formation. The retrosplenial dysgranular cortex (Rdg) projects to the postsubiculum, caudal parts of parasubiculum, caudal and lateral parts of the entorhinal cortex, and the perirhinal cortex. The retrosplenial granular b cortex (Rgb) projects only to the postsubiculum, but the retrosplenial granular a cortex (Rga) projects to the postsubiculu, rostral presubiculum, parasubiculum, and caudal medial entorhinal cortex. Reciprocating projections from the hippocampal formation to Rdg originate in septal parts of CA1, postsubiculum, and caudal parts of the entorhinal cortex, but these are only sparse projections. In contrast, Rgb and Rga receive dense projections from the hippocampal formation. The hippocampal projection to Rgb originates in area CA1, dorsal (septal) subiculum, and post-subiculum. Conversely, Rga is innervated by ventral (temporal) subiculum and postsubiculum. Further, the connections between the retrosplenial cortex and the hippocampal formation are topographically organized. Rostral retrosplenial cortex is connected primarily to the septal (rostrodorsal) hippocampal formation, while caudal parts of the retrosplenial cortex are connected with temporal (caudoventral) areas of the hippocampal formation. Together, the elaborate connections between the retrosplenial cortex and the hippocampal formation suggest that this projection provides an important pathway by which the hippocampus affects learning, memory, and emotional behavior.

Afferent Pathways↗

An anatomical study of cholinergic innervation in rat cerebral cortex.

The cholinergic innervation of rat cerebral cortex was studied by immunohistochemical localization of choline acetyltransferase. Stained bipolar cells, fibers and terminals were found in all areas of cortex. The density of cholinergic terminals was similar in all cortical areas with the exception of entorhinal and olfactory cortex, which showed a marked increase in the number of stained terminals. A laminar distribution of cholinergic terminals was found in many cortical areas. In motor and most sensory areas, terminal density was high in layer 1 and upper layer 5, and lowest in layer 4. Visual cortex, in contrast to other cortical areas, was characterized by a dense band of innervation in layer 4. It has been known that the majority of cortical cholinergic structures derive from a projection to cortex from large, multipolar neurons in the basal forebrain, which stain heavily for choline acetyltransferase. In this study, stained fibers were observed to take three different pathways from basal forebrain to cortex. The first, confined to medial aspects of forebrain and cortex, was observed to originate in the septal area, from where fibers formed a discrete bundle, swinging forward around the rostral end of the corpus callosum, then travelling caudally in the cingulate bundle. The second was found to consist of fibers fanning out laterally from the area of the globus pallidus, travelling through the caudate, then continuing for various distances in the corpus callosum before finally turning into the cortex. A third pathway appeared to innervate olfactory and entorhinal cortex. Ibotenic acid injections were made in the area of the globus pallidus to study the effect of lesioning the lateral pathway on the cholinergic innervation in cortex. A major loss of choline acetyltransferase positive terminals was observed in neocortex, but retrosplenial, cingulate, entorhinal and olfactory cortex showed a normal density of cholinergic innervation. The borders separating areas with lesioned cholinergic input from non-lesioned areas were precise. The distribution of stained terminals remaining in cortical areas with lesioned basal forebrain innervation suggests that the basal forebrain projection to cerebral cortex, and not the intrinsic cortical cholinergic neurons, give rise to the laminar distribution of cholinergic terminals observed in normal cortex. To compare the relative densities of different cholinergic cortical systems, the distribution of choline acetyltransferase staining was compared with that of vasoactive intestinal polypeptide and substance P, which are co-localized in some choline acetyltransferase-positive neurons innervating cortex.

Animals↗

Long-lasting potentiation in the secondary somatosensory cortex affects motor control: assessment by H-reflex.

We investigated descending projections from the secondary somatosensory cortex to the feline spinal cord and the effects of long-lasting potentiation in secondary somatosensory cortex on the activities of motoneurons of the cat. Electrophysiological examinations revealed that the low-intensity subthreshold secondary somatosensory cortex stimulation could change the H-Reflex induced by radial nerve stimulation. The H-wave amplitudes, recorded in wrist flexor muscles, were enhanced when the intervals from secondary somatosensory cortex to radial nerve stimuli were altered from 0 to 30 ms (initial excitation, 146 +/- 11% (mean +/- S.E.M.) of the control value). In contrast, the H-waves were suppressed with intervals longer than 30 ms (80 +/- 3%). The descending pathways from secondary somatosensory cortex to the spinal cord were assessed using an immunohistochemical technique. c-Fos and Zif268 proteins, induced by stimulation of the hand-represented secondary somatosensory cortex areas, could thus express in activated cervical neurons. The density of labeled cells was significantly higher in the seventh and eighth cervical segments than in other levels. The great majority of positive cells were distributed in the lateral part of the contralateral ventral horn and their somas ranged from 10 to 50 microns in size. Finally, we examined the effects of long-lasting potentiation, induced by high-frequency stimulation of the ventral posterolateral thalamic nucleus, on the activities of spinal motoneurons. Long-lasting potentiation altered the previously observed effects of secondary somatosensory cortex stimulation on the H-wave amplitude. The secondary somatosensory cortex-conditioned initial excitation of the H-reflex was enhanced (from 139 to 175%, P < 0.05), while late suppression was completely blocked (from 74 to 112%, P < 0.01). In conclusion, the descending pathways from secondary somatosensory cortex to the spinal cord modulated the H-reflex, and long-lasting potentiation in secondary somatosensory cortex affected this modulation. We have previously reported that corticocortical inputs from primary to secondary somatosensory cortex is required for induction of long-lasting potentiation in secondary somatosensory cortex. Taken together, the present study suggests that cortical plasticity in secondary somatosensory cortex amplifies somatic inputs from primary somatosensory cortex as a means of adaptive motor control by the sensory system.

Animals↗

Laminar distribution of nicotinic receptor subtypes in human cerebral cortex as determined by [3H](-)nicotine, [3H]cytisine and [3H]epibatidine in vitro autoradiography.

The subregional localization of different nicotinic acetylcholine receptor subtypes in human cerebral cortex was estimated by quantitative in vitro autoradiography using the nicotinic ligands [3H](-)nicotine, [3H]cytisine and [3H]epibatidine in large whole human forebrain hemispheres. Saturation experiments in frontal cortex revealed for [3H](-)nicotine two binding sites with affinity constants (Kd) of 0.45 and 6.3 nM and binding site densities (Bmax) of 3.0 and 14.2 pmol/g, for [3H]cytisine one binding site with Kd of 0.19 nM and Bmax of 21.8 pmol/g, and for [3H]epibatidine one binding site with Kd of 0.011 nM and Bmax of 20.0 pmol/g. The laminar binding distributions of the three ligands were compared in different cortical areas by creating binding profiles perpendicular to the entire cortical depth. The regional autoradiographic binding patterns of the three ligands were essentially similar, with higher receptor binding in cortical layers I, III and V. In the primary sensory cortex and inferior frontal sulcus, marked binding of all ligands was observed in layer III. [3H]Cytisine showed the lowest difference between maximal and minimal binding within the gray tissue in all other areas. In the primary motor cortex, [3H]epibatidine and [3H](-)nicotine showed high binding in layers III and V. The [3H](-)nicotine binding was higher than that of the other ligands in layers I and VI of the primary motor cortex, the deeper layer V of the primary sensory cortex, layer III of the superior temporal sulcus and layer VI of the parietal cortex. A distinct band of binding of [3H](-)nicotine and [3H]epibatidine but not of [3H]cytisine was found in layer IIlb of the occipital cortex and layer V of the superior temporal sulcus. [3H]Epibatidine showed higher binding than the other ligands in all layers of the medial frontal, superior frontal and superior temporal sulcus. The findings with the three nicotinic ligands suggest three binding sites in the cortex with different laminar distributions. All three ligands bound to an identical receptor site, most likely the alpha4 nicotinic receptor subunit. The morphological distribution of [3H]epibatidine and [3H](-)nicotine binding indicate that they bind to an additional site, especially in the primary motor cortex, in layer IIIb of the occipital cortex and layer V of the superior temporal sulcus. High binding of [3H](-)nicotine in layers I and VI of the primary motor cortex, the deeper layer V of the primary sensory cortex, layer III of the superior temporal sulcus and layer VI of the parietal cortex may indicate a third binding site.

Aged↗

The cortical projections of foveal striate cortex in the rhesus monkey.

1. The cortical projections of the foveal and extrafoveal parts of the striate cortex have been compared, using conventional degeneration techniques, as well as combinations of anatomical methods. While both foveal and extrafoveal striate cortex share a common pattern of projections (to areas V2, V3 and the visual area in the medial part of the posterior bank of the superior temporal sulcus), foveal striate cortex was found to have an additional projection (to part of the cortex of the fourth visual areas, V4). The latter projection includes the posterior lip of the inferior occipital sulcus which, on anatomical grounds, is regarded as the ventral extension of V4. 2. Anatomical studies using double tracers were employed to clarify the nature of the projections from the striate cortex and from V2 to V4. In one such experiment, tritiated proline was injected into extra-foveal striate cortex and a small lesion was made in that part of V2 receiving a direct projection from the region of the striate cortex into which the radioactive tracer was injected. Only degenerating fibres (due to the lesion), and no radioactive label, was found in V4. Such an experiment showed that, unlike foveal striate cortex, the projections from extrafoveal striate cortex to V4 are not direct, but through V2. 3. In another type of anatomical experiment using double tracers, the corpus callosum was sectioned and tritiated proline was injected into foveal striate cortex. Such an experiment allowed a more accurate determination of the extent of V4, as judged from its callosal connexions, to which foveal striate cortex projects. 4. Considering the projections of V1 to areas V2, V3 and the visual area in the medial part of the posterior bank of the superior temporal sulcus, and considering the differences in the projections of foveal and extrafoveal striate cortex, it is suggested that, among other functions, the striate cortex acts as a distribution centre for the information coming over the retino-geniculo-cortical pathways, parcelling this information out to different visual areas of the prestriate cortex for further analysis.

Animals↗

Posterior parietal cortex in rhesus monkey: I. Parcellation of areas based on distinctive limbic and sensory corticocortical connections.

Injections of HRP-WGA in four cytoarchitectonic subdivisions of the posterior parietal cortex in rhesus monkeys allowed us to examine the major limbic and sensory afferent and efferent connections of each area. Area 7a (the caudal part of the posterior parietal lobe) is reciprocally interconnected with multiple visual-related areas: the superior temporal polysensory area (STP) in the upper bank of the superior temporal sulcus (STS), visual motion areas in the upper bank of STS, the dorsal prelunate gyrus, and portions of V2 and the parieto-occipital (PO) area. Area 7a is also heavily interconnected with limbic areas: the ventral posterior cingulate cortex, agranular retrosplenial cortex, caudomedial lobule, the parahippocampal gyrus, and the presubiculum. By contrast, the adjacent subdivision, area 7ip (within the posterior bank of the intraparietal sulcus), has few limbic connections but projects to and receives projections from widespread visual areas different than those that are connected with area 7a: the ventral bank and fundus of the STS including part of the STP cortex and the inferotemporal cortex (IT), areas MT (middle temporal) and possibly MTp (MT peripheral) and FST (fundal superior temporal) and portions of V2, V3v, V3d, V3A, V4, PO, and the inferior temporal (IT) convexity cortex. The connections between posterior parietal areas and visual areas located on the medial surface of the occipital and parieto-occipital cortex, containing peripheral representations of the visual field (V2, V3, PO), represent a major previously unrecognized source of visual inputs to the parietal association cortex. Area 7b (the rostral part of the posterior parietal lobe) was distinctive among parietal areas in its selective association with somatosensory-related areas: S1, S2, 5, the vestibular cortex, the insular cortex, and the supplementary somatosensory area (SSA). Like 7ip, area 7b had few limbic associations. Area 7m (on the medial posterior parietal cortex) has its own topographically distinct connections with the limbic (the posterior ventral bank of the cingulate sulcus, granular retrosplenial cortex, and presubiculum), visual (V2, PO, and the visual motion cortex in the upper bank of the STS), and somatosensory (SSA, and area 5) cortical areas. Each parietal subdivision is extensively interconnected with areas of the contralateral hemisphere, including both the homotopic cortex and widespread heterotopic areas. Indeed, each area is interconnected with as many areas of the contralateral hemisphere as it is within the ipsilateral one, though less intensively. This pattern of distribution allows for a remarkable degree of interhemispheric integration.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Afferent projections to the cingulate cortex in albino rats: a study with a retrograde labeling method using fluoro-gold.

We studied the neuronal populations that project their axons to the cingulate cortex in albino rats using the retrograde fluorescent dye of 4% Fluoro-Gold injected into the anterior, middle and posterior portions of the cingulate cortex. The result showed that the following ipsilateral structures are sending fibers to these three portions: the prefrontal cortex, frontoparietal motor cortex, indusium griseum, dorsal endopiriform nucleus, lateral part of medial mammillary nucleus, nuclei of diagonal band of Broca, anterior pretectum, anterior part of caudate-putamen, hippocampal formation, anteroventral, anteromedial, lateroposterior, ventroposterior and dorsomedial thalamic nuclei. The anterior portion of the cingulate cortex receives inputs from the following ipsi- and contralateral structures: the accessory olfactory bulbs, anterior olfactory nuclei, middle and posterior portions of the cingulate cortex. The middle cingulate cortex receives fibers only from the ipsilateral dorsal part of the lateral septal nucleus in addition from the ipsi- and contralateral anterior and posterior portions of the cingulate cortex. While the posterior portion of the cingulate cortex receives separate inputs from the following ipsilateral structures: anterodorsal thalamic nucleus, temporal cortex, entorhinal cortex, areas 17 and 18, as well as from the ipsi- and contralateral anterior and middle portions of the cingulate cortex. The present study shows that the cingulate cortex receives various kinds of inputs from the other parts of the brain which are involved in emotion, memory, vision and motion, and also suggests that there are differences in afferent projections among the anterior, middle and posterior portions of the cingulate cortex.

Afferent Pathways↗