Self-stimulation of the prefrontal cortex after microinjections of pimozide and GABA into the prefrontal cortex or ventrotegmental area in the rat.
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To study the distribution of the cortical and thalamic efferent projections from the parahippocampal cortex in the cat, a series of injections of anterogradely transported radioactively labeled amino acids were placed in different parts of the entorhinal and perirhinal cortices. Subsequently, some of the identified cortical and thalamic target areas were injected with retrograde tracers such as wheat germ-agglutinin conjugated with horseradish peroxidase (WGA-HRP) or with a fluorescent tracer--fast blue or nuclear yellow--in order to disclose the laminar origin of the parahippocampal efferent projections. The results indicate that the parahippocampal cortex gives rise to widespread projections to the association cortex, and, to a lesser extent, sends fibers to the limbic cortex and the primary sensory cortex. These projections arise mainly from the deep layers of the parahippocampal cortex and terminate predominantly in superficial layers of the cortex, with a preference for layer I. Within the cortical projections a medial-to-lateral topography could be observed such that the entorhinal cortex projects predominantly to the allocortical and periallocortical limbic areas, including parts of the subicular complex, the ventral retrosplenial and the infralimbic cortices, and olfactory related areas--i.e., the olfactory bulb, the anterior olfactory nucleus, the prepiriform cortex, and the ventral tenia tecta. The more lateral parts of the parahippocampal cortex, which surround the posterior rhinal sulcus, project in addition to extensive parts of the paralimbic association cortex that include the proisocortical cingular, prelimbic, orbitofrontal, and agranular and granular insular cortices. The most lateral portion of the parahippocampal cortex, the perirhinal cortex, furthermore issues projections to widespread neocortical areas on the lateral and medial aspects of the hemisphere that constitute part of the parasensory association cortex. Weak-to-moderate projections are found to the cortex of the middle suprasylvian and anterior ectosylvian sulci, as well as the cruciate and splenial sulci, all of which have been reported to constitute sensory convergence areas. The most marked projections from the perirhinal cortex reach a zone of neocortex directly lateral to the perirhinal cortex including ventral parts of the posterior sylvian, posterior ectosylvian, posterior suprasylvian, and lateral gyri. These projections appear to be topographically organized such that rostral parts of the perirhinal cortex project more rostrally, and more caudal parts of the perirhinal cortex project to more caudal parts of this cortical zone.(ABSTRACT TRUNCATED AT 400 WORDS)
An influence of the prefrontal cortex on noradrenergic locus coeruleus neurons would have profound implications for the function of the locus coeruleus system. Although the medial prefrontal cortex does not substantially innervate the core of the nucleus locus coeruleus, evidence indicates that the medial prefrontal cortex projects to regions containing locus coeruleus dendrites; indirect medial prefrontal cortex-locus coeruleus projections are also possible. Here, we examined influences of prefrontal cortex activity on locus coeruleus firing rates by activating or inactivating the medial prefrontal cortex while recording impulse activity of locus coeruleus neurons extracellularly in anaesthetized rats. Most of our electrical stimulation experiments were conducted in rats which underwent lesions of the ascending dorsal bundle of noradrenergic fibres from the locus coeruleus to eliminate locus coeruleus projections to the prefrontal cortex, because antidromic activation of locus coeruleus from the prefrontal cortex affects even non-driven locus coeruleus neurons through collaterals. Single pulse stimulation (1 mA, 0.3-0.5 ms) of the dorsomedial (frontal region 2) or prelimbic region of the medial prefrontal cortex synaptically activated 13/16 (81%) or 16/56 (29%) locus coeruleus neurons, respectively. Train stimulation (20 Hz for 0.5 s) synaptically activated greater percentages of locus coeruleus cells, 11/12 cells (92%) for the dorsomedial prefrontal cortex, and 41/50 cells (82%) for the prelimbic cortex. No inhibitory responses in the locus coeruleus were obtained with dorsomedial prefrontal stimulation, and weak inhibition was found in 16% of locus coeruleus cells with prelimbic stimulation. Electrical stimulation of more lateral frontal cortex (Fr1 area) had no effects on locus coeruleus activity. Chemical stimulation of the dorsomedial prefrontal cortex with L-glutamate (10 or 100 mM) or D,L-homocysteic acid (10 mM) phasically activated 15/26 (55%) locus coeruleus cells, and 15/68 cells (22%) with prelimbic stimulation; such activation was sometimes followed by long-lasting oscillatory activity. No locus coeruleus cells exhibited purely inhibitory responses with chemical stimulation of any prefrontal cortex site. Inactivation of the dorsomedial or prelimbic region of the prefrontal cortex with lidocaine microinjection (2%, 180 or 300 nl) reduced locus coeruleus firing rates in 6/10 (60%) or 7/19 (37%) locus coeruleus cells, respectively. In no case did lidocaine in any prefrontal cortex site activate a locus coeruleus neuron. These results indicate that the medial prefrontal cortex provides a potent excitatory influence on locus coeruleus neurons. The fact that inactivation of the medial prefrontal cortex suppressed locus coeruleus firing indicates that the medial prefrontal cortex also provides a resting tonic excitatory influence on locus coeruleus activity.
Thalamic and cortical afferents to the insular and perirhinal cortex of the rat were investigated. Unilateral injections of horseradish peroxidase (HRP) were made iontophoretically along the rhinal sulcus. HRP injections covered or invaded areas along the rhinal fissure from about the level of the middle cerebral artery to the posterior end of the fissure. The most anterior injection labeled a few cells in the mediodorsal nucleus. More posterior injections labeled neurons in the basal portion of the nucleus ventralis medialis, thus suggesting that this cortical region constitutes the rat's gustatory (insular) cortex. We consider the cortex situated posterior to the gustatory cortex in and above the rhinal sulcus as the core region of the rat's (associative) insular cortex, as this cortex receives afferents from the regions of and between the nuclei suprageniculatus and geniculatus medialis, pars magnocellularis. It includes parts of the cortex termed perirhinal in other studies. The cortex dorsal and posterior to the insular cortex we consider auditory cortex, as it receives afferents from the principal part of the medial geniculate nucleus, and the cortex ventral to the insular cortex (below the fundus of the rhinal sulcus) we consider to constitute the prepiriform cortex, which is athalamic. The posterior part of the perirhinal cortex (area 35) receives afferents from nonspecific thalamic nuclei (midline nuclei). Cortical afferents to the injection loci arise from a number of regions, above all from regions of the medial and sulcal prefrontal cortex. Those injections confined to the projection cortex of the suprageniculate-magnocellular medial geniculate nuclear complex also led to labeling in contralateral prefrontal regions, particularly in area 25 (infralimbic region). A comparison of our results with those on the insular cortex of cats and monkeys suggests that on the basis of thalamocortical connections, topographical relations, and involvements of neurons in information processing and overt behavior, the insular cortex has to be regarded as a heterogeneous region which may be separated into prefrontal insular, gustatory (somatosensory) insular, and associative insular portions.
Recent studies have explored certain changes with aging of neurons containing neuropeptides. The degree of loss of vasoactive intestinal polypeptide (VIP)-, neuropeptide Y (NPY)- and somatostatin-containing neurons in the aged CNS has not yet been established with certainty however, and available data is often contradictory. Changes with aging of VIP- and NPY-containing neurons were demonstrated by immunocytochemistry in this study. A major loss of VIP-immunoreactive (ir) neurons in aged rat brain was observed in the frontal cortex area 3, parietal cortex area 1, hindlimb area, temporal cortex area 1 and 2, monocular part of occipital cortex area 1, occipital cortex area 2, and retrosplenial cortex. VIP-ir cells in the frontal cortex areas 1 and 2, parietal cortex area 2, forelimb area, binocular part of the occipital cortex area 1, and the dentate gyrus were moderately decreased. The axis of VIP neurons in the aged group showed an irregular orientation tendency, especially in layers II and III. Major loss of NPY-ir neurons in aged rat brain were observed in the retrosplenial cortex, frontal cortex areas 1 and 2, parietal cortex areas 1 and 2, occipital cortex areas 1 and 2, the temporal cortex, hippocampus proper and cingulate cortex. Loss of NPY-ir neurons was observed mostly in layers V and VI. The number and length of dendritic branches also appeared to have decreased and shortened in the aged group. There were only slight decreases of somatostatin-ir cell numbers in the parietal and occipital cortex of the aged group. These results indicate the involvement of VIP and NPY-ir neurons in the aging process of cerebral cortex, and provide the morphological evidence for the decreased number of VIP and NPY neurons by immunocytochemistry in each area of cerebral cortex of aged rats.
A number of sensory modalities most likely converge in the rat perirhinal cortex. The perirhinal cortex also interconnects with the amygdala, which plays an important role in various motivational and emotional behaviors. The neural pathway from the perirhinal cortex to the entorhinal cortex is considered one of the main paths into the entorhinal-hippocampal network, which has a crucial role in memory processes. To investigate the potential associative function of the perirhinal cortex with respect to sensory and motivational stimuli and the influence of the association on the perirhinal-entorhinal-hippocampal neurocircuit, we prepared rat brain slices including the perirhinal cortex, entorhinal cortex, hippocampal formation, and amygdala. We used an optical imaging technique with a voltage-sensitive dye to analyze 1) the spatial and functional distribution of inputs from the lateral nucleus of the amygdala to the perirhinal cortex; 2) the spread of neural activity in the perirhinal cortex after layers II/III stimulation, which mimics sensory input to the perirhinal cortex; and 3) the effect of associative inputs to the perirhinal cortex from both the lateral amygdaloid nucleus and layers II/III of the perirhinal cortex on the perirhinal-entorhinal-hippocampal neurocircuit. Following stimulation in the superficial layers of the perirhinal cortex, electrical activity only propagated into the entorhinal cortex when sufficient activation occurred in the deep layers of perirhinal area 35. We observed that single stimulation of either the perirhinal cortex or amygdala did not result in sufficient neural activation of the deep layers of areas 35 to provoke activity propagation into the entorhinal cortex. However, the deep layers of area 35 were depolarized much more strongly when the two stimuli were applied simultaneously, resulting in spreading activation in the entorhinal cortex. Our observations suggest that a functional neural basis for the association of higher-order sensory inputs and emotion-related inputs exists in the perirhinal cortex and that transfer of sensory information to the entorhinal-hippocampal circuitry might be affected by the association of that information with incoming information from the amygdala.
The flow of information in the sensorimotor cortex may determine how somatic information modulates motor cortex neuronal activity during voluntary movement. Electrophysiological recordings and neuroanatomical tracing techniques were used to study the connections between the primary somatosensory cortex (SI) and the vibrissal representation of the primary motor cortex (MI) in rodents. Intracortical microstimulation (ICMS) was applied to the vibrissal region of the motor cortex to identify a site from which stimulation evoked movements of the vibrissae. Movements of only a single whisker were evoked by applying low-intensity stimulating current to particular locations within MI. A single injection of either horseradish peroxidase (HRP) or biocytin was made at the stimulus site in each animal, to retrogradely label cells in the somatosensory cortex. Receptive field (RF) responses were recorded from neurons in the barrel cortex to identify the sensory cortex representation of the same whisker that responded to ICMS. The site at which neurons responded predominately to manual stimulation of this particular vibrissa was marked by a small electrolytic lesion. The projection from the somatosensory cortex to the identified whisker representation in the motor cortex was determined by mapping the location of labeled neurons in tissue sections processed for either HRP or biocytin. The relationship of the labeled cells in SI to the barrel structures was determined from adjacent sections that were stained for cytochrome oxidase. In all cases, the barrel column associated with the relevant whisker contained labeled cells. Surrounding barrels also contained labeled cells, although fewer in number. Very few labeled cells were found in non-contiguous barrels. These results show that the SI to MI projection is somatotopically arranged, such that the sensory cortex representation of a whisker is morphologically connected to the motor cortex representation of the same whisker. Thus, sensory information is relayed to MI from the relevant whisker region in SI. Adjacent whisker regions also appear to relay somatic input, but presumably to a lesser degree. A second group of animals received single small injections of the anterograde tracer, Phaseolus vulgaris leucoagglutinin, to an electrophysiologically identified whisker representation in the sensory cortex. A single narrow column of labeled fibers was found in the motor cortex following such injections. Thus, the sensory cortex appears to relay somatic information from the vibrissae to restricted regions of the motor cortex in a somatotopically organized manner. Furthermore, the stimulus-evoked whisker movements suggest that certain features of the output map of the motor cortex are discretely organized.(ABSTRACT TRUNCATED AT 400 WORDS)
Corticocortical fiber projections of the striate and occipital cortex of the rabbit, as degined by Rose ("31), have been determined by fiber degeneration methods following the production of cortical lesions within each of 24 rabbits. We have assumed that the striate and occipital cortices correspond respectively to the visual cortical areas 1 and 2 (VI and V2) which have been demarcated electrophysiologically by Thompson et al. ("50). A study of the ipsilateral fiber projections of the striate and occipital cortex of the rabbit reveals three distinct sets of associational corticocortical connections. (1) Neurons located in layers I-III of all regions of the striate cortex and the occipital cortex send fibers to terminate prodominantly in layer V, but also in layers IV and VI, immediately beneath the cells of origin; however, the cells in the supragranular layers have not been found to send fibers to any other region of cerebral cortex. (2) The binocular portions of VI and V2 appear to be interconnected ipsilaterally since cells in layers IV-VI of the lateral striate cortex have been shown to project to all layers of a restricted, adjacent portion of the medial occipital cortex; and the cells in layers IV-VI of medial occipital cortex send a similar, restricted projection to the adjacent lateral striate cortex. (3) Nerve cells in layers IV-VI of the lateral striate cortex (binocular VI) send a restricted projection to the lateral portion of the occipital cortex. (4) After all lesions of the striate and/or occipital cortices, degenerating fibers are seen radiating away from the lesion in layer I; the origin of these degenerating fibers could not be determined. The following observations have been made concerning the origins and terminations of commissural corticortical fibers. (1) after ablation of most of the visual cortex of one side, commissural fibers are seen to terminate in all cortical layers in two narrow bands of visual cortex: one band occupies both sides of the striate-occipital boundary; the second band is found in the lateral portion of occipital cortex. (2) More punctate lesions reveal that commissural fibers arise from layers IV-VI of the lateral striate cortex and medial occipital cortex (binocular portions of V1 and V2 respectively) and end in homotopic areas of the contralateral cortex.
The cortical afferents to the cortex of the anterior ectosylvian sulcus (SEsA) were studied in the cat, using the retrograde axonal transport of horseradish peroxidase technique. Following injections of the enzyme in the cortex of both banks, fundus and both ends (postero-dorsal and anteroventral) of the anterior ectosylvian sulcus, retrograde labeling was found in: the primary, secondary, and tertiary somatosensory areas (SI, SII and SIII); the motor and premotor cortices; the primary, secondary, anterior and suprasylvian fringe auditory areas; the lateral suprasylvian (LS) area, area 20 and posterior suprasylvian visual area; the insular cortex and cortex of posterior half of the sulcus sylvius; in area 36 of the perirhinal cortex; and in the medial bank of the presylvian sulcus in the prefrontal cortex. Moreover, these connections are topographically organized. Considering the topographical distribution of the cortical afferents, three sectors may be distinguished in the cortex of the SEsA. The cortex of the rostral two-thirds of the dorsal bank. This sector receives cortical projections from areas SI, SII and SIII, and from the motor cortex. It also receives projections from the anterolateral subdivision of LS, and area 36. The cortex of the posterior third of the dorsal bank and of the posterodorsal end. It receives cortical afferents principally from the primary, secondary and anterior auditory areas, from SI, SII and fourth somatosensory area, from the anterolateral subdivision of LS, vestibular cortex and area 36. The cortex of the ventral bank and fundus. This sulcal sector receives abundant connections from visual areas (LS, 20, posterior suprasylvian, 21 and 19), principally from the lateral posterior and dorsal subdivisions of LS. It also receives abundant connections from the granular insular cortex, caudal part of the cortex of the sylvian sulcus and suprasylvian fringe. Less abundant cortical afferents were found to arise in area 36, second auditory area and prefrontal cortex. The abundant sensory input of different modalities which appears to converge in the cortex of the anterior ectosylvian sulcus, and the consistent projection from this cortex to the deep layers of the superior colliculus, make this cortical region well suited to play a role in the control of the orientation movements of the eyes and head toward different sensory stimuli.
The effects of neonatal administration of the catecholamine neurotoxin 6-hydroxydopamine (6-OHDA; 1-4 doses of 100 mg/kg body weight s.c.) on the postnatal development of pyramidal neurons in several cortical regions of the rat was studied using a Golgi-Cox neuronal impregnation technique. Rats were sacrificed in the adult stage (eight weeks) and the following regions were studied: anterior frontal cortex, posterior frontal cortex (including motor cortex), anterior parietal cortex (including sensory cortex), posterior parieto-occipital cortex and cingulate cortex. Significant alterations were seen in animals which received four doses of 6-OHDA. These alterations can be summarized as follows: (1) a decreased length and branching of basolateral dendrites of pyramidal cells, with loss of dendritic spines, which were found in both the internal pyrimidal layer (layer V) and the external pyramidal layer (layer III), most abundantly in the frontal cortex and cingulate cortex; (2) an increased number of pyramidal cells of layer V with premature apical dendritic termination in layer III rather than the usual termination in layers I and II. This was most abundant in the cingulate cortex; (3) occasional disorientation of pyramidal cell apical dendrites away from the normal vertical plane by 15 or more degrees, seen in frontal, parietal and cingulate cortex; (4) an increased number of pyramidal cells with rounded somatic contours, found in frontal, anterior parietal and cingulate cortex. These phenomena were occasionally seen in normal cortex, but were significantly increased in their occurrence after four doses of 6-OHDA. Such alterations were not significant in rats treated with one or three doses of 6-OHDA. The extent and severity of morphological alterations correlate with reductions in endogenous noradrenaline (NA) in cerebral cortex, which was found to average 50% of control levels after one dose of 6-OHDA, and 80% reduction after three doses, and a 97-98% reduction after four doses, suggesting that the NA denervation must be almost complete to result in readily detectable significant morphological changes in the development of cortical pyramidal cells. No consistent changes in endogenous dopamine (DA) levels were observed, except for an increase in the cingulate cortex. The anatomical alterations in pyramidal cells described in the present study suggest that NA neurons which project into the cerebral cortex have a neurotrophic role in the postnatal development of cortex.
The connections of rat cingulate cortex with visual, motor, and postsubicular cortices were investigated with retrograde and anterograde tracing techniques. In addition, connections between visual and the postsubicular (area 48) and parasubicular (area 49) cortices were evaluated with the same techniques. The following conclusions were drawn. Area 29 connections: Afferents to area 29 originate mainly from cingulate areas 24 and 25, visual cortex (primarily area 18b), motor cortex area 8, area 11 of frontal cortex, areas 48 and 49, and the subiculum. Efferent connections of area 29 within cingulate cortex and to visual areas differ for each cytoarchitectural subdivision of area 29. Thus, area 29c has limited projections both within cingulate cortex and to areas 48 and 49, while area 29d projects to these areas as well as to area 8, area 18b, and medial area 17. These visual cortex afferents originate mainly from layer V neurons of areas 29b and 29d, while areas 29a and 29c have virtually no projections to visual cortex. Area 24 connections: Afferents to area 24 originate primary from cingulate areas 25 and 29 and visual area 18b and medial area 17. Efferent projections of area 24a are distributed within cingulate cortex, while area 24b has more extensive projections to posterior cingulate and visual cortices. Area 24b is the cingulate subdivision which is both the primary recipient of visual cortex afferents as well as the source of most of the projections of anterior cingulate cortex to visual areas. Visual cortex has reciprocal connections with parts of the postsubicular and parasubicular cortices. Neurons of the internal pyramidal cell layer of both areas 48 and 49 project to areas 17 and 18b, while layers I and III of these parahippocampal areas receive projections from areas 17 and 8b. In conclusion, areas 29d have particularly extensive interconnections with visual cortex, while area 29d also maintains projections to area 8 of motor cortex. This connection scheme supports the view that cingulate cortex may have a role in feature extraction from the sensory environment, as well as in sensorimotor integration. Finally, the postsubiculum may be classified as a limbic association cortex in which extensive visual and cingulate efferents converge.
The association and commissural fiber systems arising in the olfactory cortical areas caudal to the olfactory peduncle (the piriform cortex, nucleus of the lateral olfactory tract, anterior cortical nucleus of the amygdala, periamygdaloid cortex and entorhinal cortex) have been studied utilizing horseradish peroxidase as both an anterograde and a retrograde axonal tracer. In the piriform cortex two sublaminae within layer II (IIa and IIb) layer III have been found to give rise to distinctly different projections. Retrograde cell labeling experiments indicate that the association fiber projection from layer IIb is predominatnly caudally directed, while the projection from layer III is predominantly rostrally directed. Cells in layer IIa project heavily to areas both caudal and rostral to the piriform cortex. The commissural fibers from the piriform cortex are largely restricted in their origin to layer IIb of the anterior part of the piriform cortex and in their termination on the contralteral side to the posterior part of the piriform cortex and adjacent olfactory cortical areas. A projection to the olfactory bulb has also been found to arise from cells in layers IIb and III of the ipsilateral piriform cortex, but not in layer IIa. In addition to those from the piriform cortex, association projections have also been found from other olfactory cortical areas. The nucleus of the lateral olfactory tract has a heavy bilateral projection to the medial part of the anterior piriform cortex and the lateral part of the olfactory tubercle (as well as a lighter projection to the olfactory bulb); both the anterior cortical nucleus of the amygdala and the periamygdaloid cortex project ipsilaterally to several olfactory cortical areas. The entorhinal cortex has been found to project to the medial parts of the olfactory tubercle and the olfactory peduncle. The olfactory tubercle is the only olfactory cortical area from which no association fiber systems (instrinsic or extrinsic) have been found to originate. A broad topographic organization exists in the distribution of the fibers from several of the olfactory areas. This is most obvious in the anterior part of the olfactory cortex, in which fibers from the more rostral areas (the anterior olfactory nucleus and the anterior piriform cortex) terminate in regions near the lateral olfactory tract, while those from more caudal areas (the posterior piriform cortex and the entorhinal cortex) terminate in areas further removed, both laterally and medially, from the tract. Projection to olfactory areas from the hypothalamus, thalamus, diagonal band, and biogenic amine cell groups have been briefly described.
The thalamic projections to somatic sensory-motor (SSM) cortex and adjacent cortical areas of the Virginia opossum were studied using anterograde and retrograde axoplasmic transport techniques. Large injections of horseradish peroxidase and/or tritiated amino acids were made in the parietal cortex to identify all of the thalamic nuclei that are interconnected with this large cortical area. Very restricted injections were then made in physiologically identified subdivisions of SSM cortex, in the remaining posterior portion of parietal cortex, and in the anteriorly adjacent postorbital cortex. The results show that the parietal cortex is reciprocally connected with a number of thalamic nuclei. Different combinations of these thalamic areas project to specific subregions within the parietal field. All parts of the SSM cortex, which occupies the anterior four-fifths of parietal cortex, receive input from the ventrobasal complex (VB), the ventrolateral complex (VL), the central intralaminar nucleus (CIN), the central lateral nucleus (CL), and the ventromedial nucleus (VM). We could detect no segregation of VL and VB inputs in any part of SSM cortex. Projections from all of these thalamic nuclei, except VM, show at least some degree of topographic organization. Anterior-posterior strips of SSM cortex receive input from clusters of thalamic neurons that extend dorsoventrally and rostrocaudally through VB and VL. The posterior one-fifth of the parietal cortex (the posterior parietal area) receives input from VL, the posterior nuclear complex, and the lateral complex, as well as input from CL, CIN, and VM. Postorbital cortex receives input mainly from intralaminar, midline, and medial thalamic nuclei. We conclude that the projection field of VB in the parietal cortex coincides precisely with the first somatic sensory area (SI) as defined by single unit studies (Pubols et al., '76). The VB projection field also delineates the area of the first motor (MI) representation. Thus, there is no separation of SI and MI cortex in the opossum. The posterior parietal area lies outside of SSM cortex and has thalamic connections similar to the posterior parts of parietal cortex in other mammals.
On cytoarchitectonic grounds we have identified two distinct types of cortical formations composing the lateral gyrus (visual cortex) of the dolphin and have termed these heterolaminar cortex and homolaminar cortex. The heterolaminar cortex occupies the medial and lateral banks of the entolateral sulcus whereas the homolaminar cortex occupies the remainder of the lateral gyrus both lateral and medial to the entolateral sulcus. Each of these cortices exhibits special cytoarchitectonic features, a major difference being that heterolaminar cortex contains an incipient layer IV whereas layer IV is clearly absent in homolaminar cortex. Quantitative imaging procedures reveal that there is greater laminar differentiation in heterolaminar than in homolaminar cortex. Golgi analysis of neuronal forms and dendritic architecture confirms this distinction between the two types of cortex composing the lateral gyrus. Computer-assisted morphometric methods have been applied to both types of cortex and indicate by a variety of parameters several quantitative differences in the cellular numbers, types, and organization in each type of cortex. Both types of cortex, homolaminar and heterolaminar, exhibit a markedly higher cellular density in the posterior sector of the lateral gyrus than in the anterior sector. We have also for the first time been able to identify a columnar type of organization of the cetacean visual cortex and have described two types of cytoarchitectonic columns, major and minor, in each of these types of cortex. Comparisons in organization of these basic columnar units between the bat, representing a prototypic brain, and the dolphin reveal many similarities but also major quantitative differences in type of organization between the visual cortices in these species. Marked differences are also seen between the cytoarchitectonic columnar organization of the visual cortices in the dolphin and columnar organization of striate cortex in the human brain, the number of columns per unit of cortex in the human being almost twice that seen in the dolphin brain. Some phylogenetic implications of these findings are discussed in relation to the so-called "initial" type of cortical organization reconstructed largely by retrospective inference.
The present report is the last in a series of papers on the connectivity of the parahippocampal cortex in the cat, which in this species is considered to be composed of the entorhinal and perirhinal cortices. Injections of anterogradely transported tritiated amino acids and the retrograde tracers HRP, WGA-HRP, fast blue, or nuclear yellow were placed within the limits of the parahippocampal cortex. An analysis was made of the resulting pattern of anterograde labeling and of the distribution of retrogradely labeled neurons within the parahippocampal cortex. It appears that within the parahippocampal cortex of the cat a framework exists, which is composed of longitudinal and transverse connections, organized according to three principles: Medially directed projections originate mostly in superficial layers, whereas laterally directed fibers come from deep layers. The longitudinal connections span the entire rostrocaudal extent of the parahippocampal cortex, whereas the mediolateral extent of the transverse connections is in general more restricted. Based on the organization of these longitudinal and transverse connections four longitudinal zones are recognized. The lateral entorhinal cortex (LEA) projects both within the entorhinal cortex and to the perirhinal cortex, whereas the intrinsic projections of the medial entorhinal cortex (MEA) are confined to the entorhinal cortex. These results are discussed in conjunction with the main organizational features of the afferent and efferent connections of the parahippocampal cortex of the cat. The premise is made that the cytoarchitectonically defined subdivisions of the cortex can be grouped into four areas, each with its own set of fiber connections and subserving different functional roles. A lateral area, constituted by the perirhinal areas 35 and 36, and the caudally adjacent postsplenial cortex, serves as a peripheral area through which the rest of the parahippocampal cortex--i.e., LEA and MEA, and ultimately the hippocampal formation--reciprocally communicates with extensive neocortical, subcortical, and thalamic regions associated with higher-order behavior. The medial part of LEA, constituted by the ventrolateral (VLEA) and ventromedial (VMEA) divisions, has reciprocal connections with the hippocampal formation and with the cortex, partly via the perirhinal cortex, and is connected with a number of subcortical structures such as the amygdala and the striatum.(ABSTRACT TRUNCATED AT 400 WORDS)
The orbitofrontal cortex of the monkey can be subdivided into a caudal agranular sector, a transitional dysgranular sector, and an anterior granular sector. The neural input into these sectors was investigated with the help of large horseradish peroxidase injections that covered the different sectors of orbitofrontal cortex. The distribution of retrograde labeling showed that the majority of the cortical projections to orbitofrontal cortex arises from a restricted set of telencephalic sources, which include prefrontal cortex, lateral, and inferomedial temporal cortex, the temporal pole, cingulate gyrus, insula, entorhinal cortex, hippocampus, amygdala, and claustrum. The posterior portion of the orbitofrontal cortex receives additional input from the piriform cortex and the anterolateral portion from gustatory, somatosensory, and premotor areas. Thalamic projections to the orbitofrontal cortex arise from midline and intralaminar nuclei, from the anteromedial nucleus, the medial dorsal nucleus, and the pulvinar nucleus. Orbitofrontal cortex also receives projections from the hypothalamus, nucleus basalis, ventral tegmental area, the raphe nuclei, the nucleus locus coeruleus, and scattered neurons of the pontomesencephalic tegmentum. The non-isocortical (agranular-dysgranular) sectors of orbitofrontal cortex receive more intense projections from the non-isocortical sectors of paralimbic areas, the hippocampus, amygdala, and midline thalamic nuclei, whereas the isocortical (granular) sector receives more intense projections from the dorsolateral prefrontal area, the granular insula, granular temporopolar cortex, posterolateral temporal cortex, and from the medial dorsal and pulvinar thalamic nuclei. Retrograde labeling within cingulate, entorhinal, and hippocampal cortices was most pronounced when the injection site extended medially into the dysgranular paraolfactory cortex of the gyrus rectus, an area that can be conceptualized as an orbitofrontal extension of the cingulate complex. These observations demonstrate that the orbitofrontal cortex has cytoarchitectonically organized projections and that it provides a convergence zone for afferents from heteromodal association and limbic areas. The diverse connections of orbitofrontal cortex are in keeping with the participation of this region in visceral, gustatory, and olfactory functions and with its importance in memory, motivation, and epileptogenesis.
The cortical regions dorsally adjacent to the posterior rhinal sulcus in the rat can be divided into a rostral region, the perirhinal cortex, which shares features of the monkey perirhinal cortex, and a caudal region, the postrhinal cortex, which has connectional attributes similar to the monkey parahippocampal cortex. We examined the connectivity among the rat perirhinal (areas 35 and 36), postrhinal, and entorhinal cortices by placing anterograde and retrograde tracers in all three regions. There is a dorsal-to-ventral cascade of connections in the perirhinal and entorhinal cortices. Dorsal area 36 projects strongly to ventral area 36, and ventral area 36 projects strongly to area 35. The return projections are substantially weaker. The cascade continues with the perirhinal to entorhinal connections. Area 35 is more strongly interconnected with the entorhinal cortex, ventral area 36 somewhat less strongly, and dorsal area 36 projects only weakly to the entorhinal cortex. The postrhinal-to-perirhinal connections also follow this general pattern. The postrhinal cortex is more heavily connected with dorsal area 36 than with ventral area 36 and is more heavily connected with area 36 than with area 35. The rostral portion of the postrhinal cortex has the strongest connections with the perirhinal cortex. Like in the monkey, the perirhinal and postrhinal cortices have different patterns of projections to the entorhinal cortex. The perirhinal cortex is preferentially connected with the rostrolateral portion of the entorhinal cortex. The postrhinal cortex projects to a part of this same region but is also connected to caudal and redial portions of the entorhinal cortex. The perirhinal and postrhinal projections to the entorhinal cortex originate in layers III and V and terminate preferentially in layers II and III.