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Age-dependent appearance of an insulo-parietal cortical signal propagation that elicits a synchronized population oscillation in the parietal cortex in rats.

We investigated postnatal development of a functional connectivity from the gustatory insular cortex to the parietal cortex, which is known to contain many more neurons responding to oral somesthetic stimulation than the insular cortex, in slices obtained from 9-35-day-old rats. Field potentials were evoked by stimulation to the insular cortex. In the mature cortex, insular stimulation elicited a solitary field potential in both the insular and parietal cortices and, as the simulation continued, the initial solitary potential came to be followed by a population oscillation of field potential in the parietal cortex, but not in the insular cortex. In the immature cortex, by contrast, insular stimulation failed to evoke both the initial solitary potential and the subsequent population oscillation in the parietal cortex. In the mature cortex, application of neither thapsigargin nor AP5 prevented elicitation of the initial solitary potential in the parietal cortex, but either of them abolished the parietal oscillation. In immature cortex bathed with low Mg(2+) medium, insular stimulation elicited both the initial solitary potential and the subsequent parietal oscillation, which were both prevented by thapsigargin or AP5. Theses results suggest that the insular and parietal cortices are anatomically connected but functionally unlinked at an early postnatal stage, and that a functional linkage, dependent both on NMDA and ryanodine receptors, is formed during the first postnatal month.

2-Amino-5-phosphonovalerate↗

Bilateral lesions of the central but not anterior or posterior parts of the piriform cortex retard amygdala kindling in rats.

The piriform cortex is thought to be involved in temporal lobe seizure propagation, such as that occurring during kindling of the amygdala or hippocampus. A number of observations suggested that the circuits of the piriform cortex might act as a critical pathway for limbic seizure discharges to assess motor systems, but direct evidence for this suggestion is scarce. Furthermore, the piriform cortex is not a homogeneous structure, which complicates studies on its role in limbic epileptogenesis. We have previously reported data indicating that the central part of the piriform cortex might be particularly involved during amygdala kindling. In order to further evaluate the role of different parts of the piriform cortex during kindling development, we bilaterally destroyed either the central, anterior or posterior piriform cortex by microinjections of ibotenate two weeks before onset of amygdala kindling. Lesions of the anterior piriform cortex hardly affected kindling acquisition, except that fewer animals exhibited stage 3 (unilateral forelimb) seizures compared to sham controls. Lesions of the central piriform cortex significantly retarded kindling, which was due to a decreased progression from stage 3 to stage 4/5 seizures, i.e. the lesioned rats needed significantly longer for the acquisition of generalized clonic seizures in the late stages of kindling development. Lesions of the posterior piriform cortex did not significantly affect kindling development. The data demonstrate that different parts of the piriform cortex mediate qualitatively different effects on amygdala kindling. The central piriform cortex seems to be a neural substrate involved in the continuous development of kindling from stage 3 to stages 4/5, indicating that this part of the piriform cortex may have preferred access, either directly or indirectly, to structures capable of supporting generalized kindled seizure expression.

Amygdala↗

Functional compensation in the lateral suprasylvian visual area following bilateral visual cortex damage in kittens.

Previous studies have shown that functional compensation is present in the cat's posteromedial lateral suprasylvian (PMLS) area of cortex after damage to areas 17, 18, and 19 (visual cortex) early in life but not after damage in adults. These studies all have investigated animals with a unilateral visual cortex lesion, whereas all behavioral studies of compensation for early visual cortex damage have investigated animals with a bilateral lesion. In the present experiment, we investigated whether functional compensation also is present in PMLS cortex after a bilateral visual cortex lesion early in life. We recorded from single neurons in the PMLS cortex of adult cats that had received a bilateral lesion of areas 17, 18, and 19 on the day of birth or at 8 weeks of age. We found that PMLS cells in both groups of cats had functional compensation (normal direction selectivity and ocular dominance) similar to that seen after a unilateral lesion at the same ages. These results are consistent with the hypothesis that PMLS cortex is involved in the behavioral compensation seen after early visual cortex damage. In addition, the results indicate that inputs from contralateral visual cortex are not necessary for the development of functional compensation seen in PMLS cortex.

Age Factors↗

Peripheral input pathways to the monkey motor cortex.

We have shown (Asanuma et al., 1979c) that the monkey motor cortex receives peripheral somesthetic inputs directly from the thalamus. In the present experiments, we studied the pathways which mediated these inputs by stimulating superficial radial (SR) and deep radial (DR) nerves and recording evoked potentials from the motor and sensory cortices and the following results were obtained: 1. The focus for SR and DR evoked potentials in the sensory cortex was located in a circumscribed small area whereas in the motor cortex, the evoked potentials were distributed in a wide area along the central sulcus including the distal forelimb area. 2. Ablation of the sensory cortex reduced the size, but neither abolished nor changed the latency of the evoked potentials in the motor cortex. 3. Section of dorsal column nearly abolished the evoked potentials in the motor cortex, but only halved their size in the sensory cortex. 4. Section of ventrolateral cervical column including the spinothalamic tract halved the size of evoked potentials in the sensory cortex, but did not change the size in the motor cortex. 5. It is concluded that direct peripheral inputs to the motor cortex are mediated primarily through the dorsal column system whereas the peripheral inputs to the sensory cortex are mediated through both dorsal column and spinothalamic tract.

Animals↗

Postnatal development of biotinylated dextran amine-labeled corpus callosum axons projecting from the visual and auditory cortices to the visual cortex of the rat.

The distribution and morphology of developing corpus callosum (CC) axons in rat visual cortex was studied by unilateral application of the in vivo anterograde tracer biotinylated dextran amine (BDA) to the visual or auditory cortex of newborns through adults. Changes in the distribution and morphology of CC axons during development were observed. Following BDA placement only in visual cortex, nearly all CC projections were to visual cortex (homotopic CC projections). At postnatal day (PND) 5-8, labeled CC axons were found throughout the contralateral visual cortex, including area 17; these CC axons could be followed from the white matter to layer I. By PND 13, few CC axons were found in medial area 17, indicating the existence of transitory CC axons in area 17 at younger ages. Morphological changes were investigated at the area 17/18a border and showed that CC axon collaterals were not formed until PND 8, and terminal arbors were not visible until PND 13; by PND 17, the adult CC-axon terminal pattern was present. At all ages, only a few heterotopic CC projections from visual to auditory cortex were found in the gray matter, although many labeled CC axons extended laterally into the white matter underlying the auditory cortex. Following BDA placement only in auditory cortex, CC projections to both auditory (homotopic CC projections) and visual (heterotopic CC projections) cortex were observed. At all ages, the homotopic CC projections were present throughout the auditory cortex, but were not distributed homogeneously; densely labeled CC axons showed a distinct columnar organization. The heterotopic CC projections were present in all visual cortical areas, including medial area 17, in significant numbers until PND 24, but were mostly eliminated by PND 28, at which time a labeling pattern similar to the adult was found. Thus, most of the heterotopic CC projections were transitory. The present study confirms the existence of transitory CC axons projecting through all layers of the visual cortex, as revealed by DiI, and extends the DiI results by showing that these transitory CC axons arise from both homotopic and heterotopic origins. Furthermore, different sources of transitory CC axons have different timetables for elimination.

Age Factors↗

Ipsilateral connections of the anterior cingulate cortex with the frontal and medial temporal cortices in the macaque monkey.

The present study was attempted to study ipsilateral corticocortical connections of the anterior part (area 24) of the cingulate cortex of the macaque monkey by means of wheat germ agglutinin-conjugated peroxidase (WGA-HRP) method. In 2 out of 4 Japanese monkeys (Macaca fuscata) that were injected with WGA-HRP into the anterior part of the cingulate cortex, the sites of injection were successfully localized within the cortical regions corresponding to areas 24a and 24b. The results obtained from these monkeys indicate that areas 24a and 24b in the anterior part of the cingulate cortex are reciprocally connected with the prefrontal, premotor, and motor cortical regions, and also with the medial temporal cortical regions. Areas 24a and 24b were strongly connected with the lateral and medial prefrontal cortices and area 6a beta of the premotor cortex, moderately with the remaining premotor cortex, and weakly with the motor cortex. In the medial temporal cortex, areas 24a and 24b were strongly connected with the prosubiculum, entorhinal cortex (area 28), and perirhinal cortex (areas 35 and 36), and weakly with areas TF and TH of the parahippocampal gyrus, throughout their rostrocaudal extent. In addition, areas 24a and 24b projected to the molecular layer of the CAI subfield of Ammon's horn and the external pyramidal layer of the presubiculum. Our findings suggest that areas 24a and 24b of the anterior cingulate cortex may constitute relays in the reciprocal pathways between the prefrontal cortex and the hippocampal, entorhinal and/or perirhinal cortical regions.

Animals↗

Contributions of anterior cingulate cortex to behaviour.

Assessments of anterior cingulate cortex in experimental animals and humans have led to unifying theories of its structural organization and contributions to mammalian behaviour. The anterior cingulate cortex forms a large region around the rostrum of the corpus callosum that is termed the anterior executive region. This region has numerous projections into motor systems, however, since these projections originate from different parts of anterior cingulate cortex and because functional studies have shown that it does not have a uniform contribution to brain functions, the anterior executive region is further subdivided into 'affect' and 'cognition' components. The affect division includes areas 25, 33 and rostral area 24, and has extensive connections with the amygdala and periaqueductal grey, and parts of it project to autonomic brainstem motor nuclei. In addition to regulating autonomic and endocrine functions, it is involved in conditioned emotional learning, vocalizations associated with expressing internal states, assessments of motivational content and assigning emotional valence to internal and external stimuli, and maternal-infant interactions. The cognition division includes caudal areas 24' and 32', the cingulate motor areas in the cingulate sulcus and nociceptive cortex. The cingulate motor areas project to the spinal cord and red nucleus and have premotor functions, while the nociceptive area is engaged in both response selection and cognitively demanding information processing. The cingulate epilepsy syndrome provides important support of experimental animal and human functional imaging studies for the role of anterior cingulate cortex in movement, affect and social behaviours. Excessive cingulate activity in cases with seizures confirmed in anterior cingulate cortex with subdural electrode recordings, can impair consciousness, alter affective state and expression, and influence skeletomotor and autonomic activity. Interictally, patients with anterior cingulate cortex epilepsy often display psychopathic or sociopathic behaviours. In other clinical examples of elevated anterior cingulate cortex activity it may contribute to tics, obsessive-compulsive behaviours, and aberrent social behaviour. Conversely, reduced cingulate activity following infarcts or surgery can contribute to behavioural disorders including akinetic mutism, diminished self-awareness and depression, motor neglect and impaired motor initiation, reduced responses to pain, and aberrent social behaviour. The role of anterior cingulate cortex in pain responsiveness is suggested by cingulumotomy results and functional imaging studies during noxious somatic stimulation. The affect division of anterior cingulate cortex modulates autonomic activity and internal emotional responses, while the cognition division is engaged in response selection associated with skeletomotor activity and responses to noxious stimuli. Overall, anterior cingulate cortex appears to play a crucial role in initiation, motivation, and goal-directed behaviours.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

The importance of being agranular: a comparative account of visual and motor cortex.

The agranular cortex is an important landmark-anatomically, as the architectural flag of mammalian motor cortex, and historically, as a spur to the development of theories of localization of function. But why, exactly, do agranularity and motor function go together? To address this question, it should be noted that not only does motor cortex lack granular layer four, it also has a relatively thinner layer three. Therefore, it is the two layers which principally constitute the ascending pathways through the sensory (granular) cortex that have regressed in motor cortex: simply stated, motor cortex does not engage in serial reprocessing of incoming sensory data. But why should a granular architecture not be demanded by the downstream relay of motor instructions through the motor cortex? The scant anatomical evidence available regarding laminar patterns suggests that the pathways from frontal and premotor areas to the primary motor cortex actually bear a greater resemblance to the descending, or feedback connections of sensory cortex that avoid the granular layer. The action of feedback connections is generally described as "modulatory" at a cellular level, or "selective" in terms of systems analysis. By contrast, ascending connections may be labelled "driving" or "instructive". Where the motor cortex uses driving inputs, they are most readily identified as sensory signals instructing the visual location of targets and the kinaesthetic state of the body. Visual signals may activate motor concepts, e.g. "mirror neurons", and the motor plan must select the appropriate muscles and forces to put the plan into action, if the decision to move is taken. This, perhaps, is why "driving" motor signals might be inappropriate-the optimal selection and its execution are conditional upon both kinaesthetic and motivational factors. The argument, summarized above, is constructed in honour of Korbinian Brodmann's centenary, and follows two of the fundamental principles of his school of thought: that uniformities in cortical structure, and development imply global conservation of some aspects of function, whereas regional variations in architecture can be used to chart the "organs" of the cortex, and perhaps to understand their functional differences.

Brain Mapping↗

Critical periods for functional and anatomical compensation in lateral suprasylvian visual area following removal of visual cortex in cats.

Previous experiments have found that neurons in the cat's lateral suprasylvian (LS) visual area of cortex show functional compensation following removal of visual cortical areas 17, 18, and 19 on the day of birth. Correspondingly, an enhanced retino-thalamic pathway to LS cortex develops in these cats. The present experiments investigated the critical periods for these changes. Unilateral lesions of areas 17, 18, and 19 were made in cats ranging in age from 1 day postnatal to 26 wk. When the cats were adult, single-cell recordings were made from LS cortex ipsilateral to the lesion. In addition, transneuronal autoradiographic methods were used to trace the retino-thalamic projections to LS cortex in many of the same animals. Following lesions in 18- and 26-wk-old cats, there is a marked reduction in direction-selective LS cortex cells and an increase in cells that respond best to stationary flashing stimuli. These results are similar to those following visual cortex lesions in adult cats. In contrast, the percentages of cells with these properties are normal following lesions made from 1 day to 12 wk of age. Thus the critical period for development of direction selectivity and greater responses to moving than to stationary flashing stimuli in LS cortex following a visual cortex lesion ends between 12 and 18 wk of age. Following lesions in 26-wk-old cats, there is a decrease in the percentage of cells that respond to the ipsilateral eye, which is similar to results following visual cortex lesions in adult cats. However, ocular dominance is normal following lesions made from 1 day to 18 wk of age. Thus the critical period for development of responses to the ipsilateral eye following a lesion ends between 18 and 26 wk of age. Following visual cortex lesions in 2-, 4-, or 8-wk-old cats, about 30% of the LS cortex cells display orientation selectivity to elongated slits of light. In contrast, few or no cells display this property in normal adult cats, cats with lesions made on the day of birth, or cats with lesions made at 12 wk of age or later. Thus an anomalous property develops for many LS cells, and the critical period for this property begins later (between 1 day and 2 wk) and ends earlier (between 8 and 12 wk) than those for other properties.(ABSTRACT TRUNCATED AT 400 WORDS)

Aging↗

Effects on muscle activity from microstimuli applied to somatosensory and motor cortex during voluntary movement in the monkey.

It is well known that electrical stimulation of primary somatosensory cortex (SI) evokes movements that resemble those evoked from primary motor cortex. These findings have led to the concept that SI may possess motor capabilities paralleling those of motor cortex and speculation that SI could function as a robust relay mediating motor responses from central and peripheral inputs. The purpose of this study was to rigorously examine the motor output capabilities of SI areas with the use of the techniques of spike- and stimulus-triggered averaging of electromyographic (EMG) activity in awake monkeys. Unit recordings were obtained from primary motor cortex and SI areas 3a, 3b, 1, and 2 in three rhesus monkeys. Spike-triggered averaging was used to assess the output linkage between individual cells and motoneurons of the recorded muscles. Cells in motor cortex producing postspike facilitation (PSpF) in spike-triggered averages of rectified EMG activity were designated corticomotoneuronal (CM) cells. Motor output efficacy was also assessed by applying stimuli through the microelectrode and computing stimulus-triggered averages of rectified EMG activity. One hundred seventy-one sites in motor cortex and 68 sites in SI were characterized functionally and tested for motor output effects on muscle activity. The incidence, character, and magnitude of motor output effects from SI areas were in sharp contrast to effects from CM cell sites in primary motor cortex. Of 68 SI cells tested with spike-triggered averaging, only one area 3a cell produced significant PSpF in spike-triggered averages of EMG activity. In comparison, 20 of 171 (12%) motor cortex cells tested produced significant postspike effects. Single-pulse intracortical microstimulation produced effects at all CM cell sites in motor cortex but at only 14% of SI sites. The large fraction of SI effects that was inhibitory represented yet another marked difference between CM cell sites in motor cortex and SI sites (25% vs 93%). The fact that motor output effects from SI were frequently absent or very weak and predominantly inhibitory emphasizes the differing motor capabilities of SI compared with primary motor cortex.

Animals↗

Histogenesis of ferret somatosensory cortex.

The ferret has emerged as an important animal model for the study of neocortical development. Although detailed studies of the birthdates of neurons populating the ferret visual cortex are available, the birthdates of neurons that reside in somatosensory cortex have not been determined. The current study used bromodeoxyuridine to establish when neurons inhabiting the somatosensory cortex are generated in the ferret; some animals also received injections of [3H]thymidine. In contrast to reports of neurogenesis in ferret visual cortex, most neurons populating the somatosensory cortex have been generated by birth. Although components of all somatosensory cortical layers have been produced at postnatal day 0, the layers are not distinctly formed but develop over a period of several weeks. A small number of neurons continue to be produced for a few days postnatally. The majority of cells belonging to a given layer are born over a period of approximately 3 days, although the subplate and last (layer 2) generated layer take somewhat longer. Although neurogenesis of the neocortex begins along a similar time line for visual and somatosensory cortex, the neurons populating the visual cortex lag substantially during the generation of layer 4, which takes more than 1 week for ferret visual cortex. Layer formation in ferret somatosensory cortex follows many established principles of cortical neurogenesis, such as the well-known inside-out development of cortical layers and the rostro-to-caudal progression of cell birth. In comparison with the development of ferret visual cortex, however, the generation of the somatosensory cortex occurs remarkably early and may reflect distinct differences in mechanisms of development between the two sensory areas.

Animals↗

Ontogeny of somatostatin immunoreactive neurons in the medial cerebral cortex and other cortical areas of the lizard Podarcis hispanica.

The ontogeny of somatostatin immunoreactive interneurons in the cerebral cortex of the lizard Podarcis hispanica has been studied in histological series of embryos, perinatal specimens, and adults. Somatostatin immunoreactive interneurons appear in the early stages of lizard cerebral cortex ontogeny, their number increases during embryonary development, reaches a peak in early postnatal life, and decreases in adult lizards. The first somatostatin immunoreactive somata in the lizard forebrain appeared on E36, and they were located in non cortical areas. Then, on E39 and later, somatostatin immunoreactive neurons were seen in the lizard cortex in a rostral-to-caudal spatial gradient, which parallels that of the normal histogenesis of the lizard cerebral cortex. On E39, labelled somata were seen in the medial and dorsal cortex inner plexiform layers; immunoreactive puncta and dendritic processes were detectable in the inner plexiform layer of the medial cortex. On E40, labelled neurons were observed in the inner plexiform layer of the lateral cortex; labelled processes were found in the inner plexiform layers (dorsomedial, dorsal, and lateral cortices) and the outer plexiform layers (medial and dorsomedial cortices). At hatching (P0), some somatostatin immunoreactive neurons populated the external plexiform layer of the dorsomedial cortex. On P28, groups of labelled neurons appeared in the cell layer of dorsal and lateral cortices, reaching the adult-mature pattern of somatostatin immunoreactivity in the lizard cerebral cortex, i.e., labelled somata and dendritic processes populating the inner plexiform layers in addition to an axonic labelled plexus in the outermost part of the outer plexiform layers. Immunoreactive somata and processes occupied all the cortical areas, but they were especially abundant in the dorsomedial cortex. Proliferating Cell Nuclear Antigen (PCNA) immunostaining in the same histological series revealed that the number of PCNA immunoreactive nuclei in the subjacent proliferative neuroepithelium followed an inverse-complementary evolution to somatostatin, suggesting some temporal relationship between somatostatin immunoreactive cells and neurogenesis in the lizard cerebral cortex.

Animals↗

Entorhinal cortex of the rat: organization of intrinsic connections.

Two sets of experiments were carried out to examine the organization of associational connections within the rat entorhinal cortex. First, a comprehensive analysis of the areal and laminar distribution of intrinsic projections was performed by using the anterograde tracers Phaseolus vulgaris-leuocoagglutinin (PHA-L) and biotinylated dextran amine (BDA). Second, retrograde tracers were injected into the dentate gyrus and PHA-L and BDA were injected into the entorhinal cortex to determine the extent to which entorhinal neurons that project to different septotemporal levels of the dentate gyrus are linked by intrinsic connections. The regional distribution of intrinsic projections within the entorhinal cortex was related to the location of the cells of origin along the mediolateral axis of the entorhinal cortex. Cells located in the lateral regions of the entorhinal cortex gave rise to intrinsic connections that largely remained within the lateral reaches of the entorhinal cortex, i.e., within the rostrocaudally situated entorhinal band of cells that projected to septal levels of the dentate gyrus. Cells located in the medial regions of the entorhinal cortex gave rise to intrinsic projections confined to the medial portion of the entorhinal cortex. Injections made into mid-mediolateral regions of the entorhinal cortex mainly gave rise to projections to mid-mediolateral levels, although some fibers did enter either lateral or medial portions of the entorhinal cortex. These patterns were the same regardless of whether the projections originated from the superficial (II-III) or deep (V-VI) layers of the entorhinal cortex. This organizational scheme indicates, and our combined retrograde/anterograde labeling studies confirmed, that laterally situated entorhinal neurons that project to septal levels of the dentate gyrus are not in direct communication with neurons projecting to the temporal portions of the dentate gyrus. These results suggest that entorhinal intrinsic connections allow for both integration (within a band) and segregation (across bands) of entorhinal cortical information processing.

Amidines↗

Cyto- and chemoarchitecture of the cerebral cortex of an echidna (Tachyglossus aculeatus). II. Laminar organization and synaptic density.

We have examined the distribution and morphology of neurons immunoreactive for nonphosphorylated neurofilament protein (SMI-32 antibody), calcium-binding proteins (parvalbumin, calbindin, calretinin), and neuropeptide Y as well as neurons reactive for NADPH diaphorase in the cerebral cortex of the Australian short-beaked echidna (Tachyglossus aculeatus). We have also studied synaptic morphology and density in S1 somatosensory cortex and assessed parameters associated with metabolic activity of the cerebral cortex (vessel volume density, mitochondrial volume density, and mitochondrial numerical density) in semi- and ultrathin sections. SMI-32 immunoreactivity was found mostly in layer V pyramidal neurons in selected cortical regions (S1, PV, V1, A). These neurons often showed atypical morphology compared with therian cortex. Neurons immunoreactive for calcium-binding proteins were broadly similar in both morphology and distribution to those seen in therian cortex, although calretinin-immunoreactive neurons were rare. Both Gray type I and Gray type II synapses could be identified in echidna S1 cortex and were similar to those seen in therian cortex. Peak synaptic density was in upper layer IV, followed by layer I, lower layer II, and upper layer III. Most synapses were of type I (72%), although types I and II were encountered with similar frequency in lower layer II and upper layer III. The capillary volume fraction values obtained for the echidna (from 1.18% in V1 to 1.34% in S1 cortex) fall within the values for rodent cortex. Similarly, values for mitochondrial volume fraction in echidna somatosensory cortex (4.68% +/- 1.76%) were comparable to those in eutherian cortex.

Animals↗

Retrograde tracing of the subset of afferent connections in mouse barrel cortex provided by zincergic neurons.

The barrel cortex of rodents is densely innervated by a prominent subclass of glutamatergic neurons that sequester and release zinc from their synaptic boutons. These neurons may play an important role in barrel cortex function and plasticity, as zinc has been shown to modulate synaptic function by regulating neurotransmitter release, excitatory and inhibitory amino acid receptors, and second messenger signaling cascades. Here, we utilized intracortical infusions of sodium selenite to identify the source of the zincergic innervation to the mouse barrel cortex. Our results demonstrate that the majority of zincergic projections to the barrel cortex arose from ipsilateral and callosal neurons, situated in cortical layers 2/3 and 6. Regionally, these labeled neurons were most abundant within the barrel cortex itself, posterior parietal association cortex, secondary somatosensory cortex, and motor cortex. Labeled neurons were also found in other somatosensory regions corresponding to the trunk, fore- and hindlimb, as well as more distant regions such as the visual, rhinal, dorsal peduncular and insular cortices, the claustrum, and lateral and basolateral amygdaloid nuclei. Further, some mice were injected with the retrograde tracer cholera toxin subunit B to compare retrograde labeling of zincergic neurons with that of the general population of neurons innervating the barrel cortex. Our data indicate that all cortical regions providing inputs to the barrel cortex possess a zincergic component, whereas those from thalamic or brainstem structures do not. These findings demonstrate that zincergic pathways comprise a chemospecific associational network that reciprocally interconnects the barrel cortex with other cortical and limbic structures.

Animals↗

Cortical and subcortical projections of the middle temporal area (MT) and adjacent cortex in galagos.

Projections of the middle temporal visual area, MT, and of visual cortex adjoining MT were investigated with autoradiographic methods in the prosimian primate, Galago senegalensis. Ipsilateral cortical targets of MT included area 17, area 18, cortex caudal to MT, cortex ventral to MT, and parietal-occipital cortex dorsal to MT. This pattern of projections suggests that extrastriate cortex contains a number of visual subdivisions in addition to MT. Contralateral projections were to MT and parietal-occipital cortex. Projections from MT to areas 17 and 18 connected regions representing similar parts of the visual hemifield while the location of callosal projections in MT matched the location of the injection site in the other hemisphere. Label in area 17 was concentrated in layers I, III, and VI whereas other cortical areas were most densely labeled in the granular and supragranular layers. Subcortical projections of MT included the reticular nucleus of the thalamus, the lateral posterior nucleus, the superior pulvinar, the inferior pulvinar, the superior colliculus, and the pontine nuclei. The projection pattern to the superior and inferior pulvinar nuclei suggests that MT projects in a topographic manner to two subdivisions within each of these structures. Injections in cortex just outside of MT labeled area 18, inferotemporal cortex, parietal-occipital cortex, and, to a lesser extent, MT. The projections to inferotemporal cortex clearly distinguish the bordering cortex from MT. Contralateral cortical terminations were in locations corresponding to the injection site. Subcortical targets were generally similar to those seen after MT injections, although additional projections were observed depending on the location of the injection. Comparison of these results from the prosimian galago with studies in New and Old World monkeys indicates there are substantial similarities in projections. Thus, some of the cortical and thalamic subdivisions described for monkeys appear to exist in prosimians.

Animals↗

Connections of the parahippocampal cortex. I. Cortical afferents.

In the present study in the cat the parahippocampal cortex denotes the caudoventral part of the limbic lobe and is composed of the entorhinal and perirhinal cortices. The cytoarchitecture of these areas and their borders with adjacent cortical areas are briefly discussed. The organization of the cortical afferents of the parahippocampal cortex was studied with the aid of retrograde and anterograde tracing techniques. In order to identify the source of cortical afferents, injections of retrograde tracers such as wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP), or the fluorescent substances fast blue or nuclear yellow, were placed in different parts of the parahippocampal cortex. In an attempt to further disclose the topographical and laminar organization of the afferent pathways, injections of tritiated amino acids were placed in cortical areas that were found to project to the parahippocampal cortex. The results of these experiments indicate that fibers from olfactory-related areas, the hippocampus, and other parts of the limbic cortex project only to the entorhinal cortex. The afferents from olfactory structures terminate predominantly superficially, whereas hippocampal and limbic cortical afferents are directed mainly to layers deep to the lamina dissecans. Paralimbic areas, including the anterior cingulate and the prelimbic cortices on the medial aspect, and the orbitofrontal and granular and agranular insular cortices on the lateral aspect of the hemisphere, project to the entorhinal cortex and medial parts of area 35 of the perirhinal cortex. These mostly mesocortical afferents terminate in both the superficial and deep layers of the entorhinal and perirhinal cortices. Parasensory association areas, which form part of the neocortex, do not project farther medially in the parahippocampal cortex than the perirhinal areas 35 and 36. These afferents mainly stem from a rather wide rim of neocortex that lies directly adjacent to area 36 and extends from the posterior sylvian gyrus via the posterior ectosylvian gyrus into the posterior suprasylvian gyrus. There is a rostrocaudal topographical arrangement in these projections such that rostral cortical areas distribute more rostrally and caudal parts project to more caudal parts of the perirhinal cortex. The cortex of the posterior suprasylvian gyrus contains the paravisual areas 20 and 21. The posterior sylvian gyrus most probably represents a para-auditory association area, whereas the most ventral part of the posterior ectosylvian gyrus may constitute a convergence area for visual and auditory inputs.(ABSTRACT TRUNCATED AT 400 WORDS)

Amidines↗

Evidence for a viscerotopic sensory representation in the cortex and thalamus in the rat.

The functional organization of the insular cortex was studied by recording neuronal responses to visceral sensory stimuli. Horseradish peroxidase (HRP) was then iontophoresed at the recording sites to identify afferents from the ventrobasal thalamus to specific visceroceptive sites in the insular cortex. The relationship of the ventrobasal thalamus to the insular cortex and to brainstem relay nuclei for the ascending visceral projections was then examined by using the axonal transport of HRP, wheat germ agglutinin conjugated to HRP (WGA-HRP), and fluorescent dyes. Of a total of 55 neurons that were tested for responses to visceral sensory stimuli, 33 units responded to at least one visceral sensory modality: 6 received gastric mechanoreceptor input, 8 responded to taste inputs, 13 were activated by arterial chemoreceptors and/or showed respiratory related activity, and 6 responded to cardiovascular baroreceptor stimulation. On the basis of its cytoarchitecture and connections with the thalamus, the insular cortex was divided into a dorsal granular area, an intermediate dysgranular region, and a ventral agranular strip. Taste-responsive neurons were located anteriorly, primarily in the dysgranular region, whereas unit responses to general visceral modalities were distributed dorsally and posteriorly in the granular insular cortex. Gastric mechanoreceptor-responsive units were situated more dorsally and anteriorly in the granular insular cortex, while cardiopulmonary inputs were located more ventrally and posteriorly. Injections of HRP into the gustatory insular cortex resulted in retrograde labeling of neurons in the parvicellular part of the ventroposterior medial thalamic nucleus (VPMpc). Injections into the general visceral insular cortex retrogradely labeled neurons lateral to VPMpc in the ventroposterior lateral parvicellular thalamic nucleus (VPLpc). Injections of HRP, WGA-HRP, and fluorescent dyes into VPMpc and VPLpc verified that their projection to the insular cortex is topographically organized. In the same experiments, retrogradely labeled neurons in the parabrachial nucleus identified the likely subnuclei within this nucleus for relay of visceral sensory information to the thalamus. Injections of WGA-HRP into the parabrachial nucleus demonstrated that its projection to the ventrobasal thalamus is also topographically organized. These results demonstrate the relationship of general visceral and special visceral (taste) representations in the insular cortex. The ascending pathway for visceral sensory information appears to be viscerotopically organized at all levels of the neuraxis, including the insular cortex.

Action Potentials↗