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Connections of somatosensory cortex in megachiropteran bats: the evolution of cortical fields in mammals.

The cortical connections of the primary somatosensory area (SI or 3b), a caudal somatosensory field (area 1/2), the second somatosensory area (SII), the parietal ventral area (PV), the ventral somatosensory area (VS), and the lateral parietal area (LP) were investigated in grey headed flying foxes by injecting anatomical tracers into electrophysiologically identified locations in these fields. The receptive fields for clusters of neurons were mapped with sufficient density for injection sites to be related to the boundaries of fields, and to representations of specific body parts within the fields. In all cases, cortex was flattened and sectioned parallel to the cortical surface. Sections were stained for myelin and architectonic features of cortex were related to physiological mapping and connection patterns. We found patterns of topographic and nontopographic connections between 3b and adjacent anterior parietal fields 3a and 1/2, and fields caudolateral to 3b (SII and PV). Area 1/2 had both topographic and nontopographic connections with 3b, PP, and SII. Connections of SII and PV with areas 3b, 3a, and 1/2 were roughly topographic, although there was clear evidence for nontopographic connections between these fields. SII was most densely connected with area 1/2, while PV was most densely connected with 3b. SII had additional connections with fields in lateral parietal cortex and with subdivisions of motor cortex. Other connections of PV were with subdivisions of motor cortex and pyriform cortex. Laminar differences in connection patterns of SII and PV with surrounding cortex were also observed. Injections in the ventral somatosensory area revealed connections with SII, PV, area 1/2, auditory cortex, entorhinal cortex, and pyriform cortex. Finally, the lateral parietal field had very dense connections with posterior parietal cortex, caudal temporal cortex, and with subdivisions of motor cortex. Our results indicate that the 3b region is not homogeneous, but is composed of myelin dense and light regions, associated with 3b proper and invaginations of area 1/2, respectively. Connections of myelin dense 3b were different from invaginating portions of myelin light area 1/2. Our findings that 3b is densely interconnected with PV and moderately to lightly interconnected with SII supports the notion that SII and PV have been confused across mammals and across studies. Our connectional evidence provides further support for our hypothesis that area 1/2 is partially incorporated in 3b and has led to theories of the evolution of cortical fields in mammals.

Animals↗

Differential spinal projections from the forelimb areas of the rostral and caudal subregions of primary motor cortex in the cat.

We used anterograde transport of WGA-HRP to examine the topography of corticospinal projections from the forelimb areas within the rostral and caudal motor cortex subregions in the cat. We compared the pattern of these projections with those from the somatic sensory cortex. The principal finding of this study was that the laminar distribution of projections to the contralateral gray matter from the two motor cortex subregions was different. The rostral motor cortex projected preferentially to laminae VI-VIII, whereas caudal motor cortex projected primarily to laminae IV-VI. Confirming earlier findings, somatic sensory cortex projected predominantly to laminae I-VI inclusive. We found that only rostral motor cortex projected to territories in the rostral cervical cord containing propriospinal neurons of cervical spinal segments C3-4 and, in the cervical enlargement, to portions presumed to contain Ia inhibitory interneurons. We generated contour maps of labeling probability on averaged segmental distributions of anterograde labeling for all analyzed sections using the same algorithm. For rostral motor cortex, heaviest label in the dorsal part of lamina VII in the contralateral cord was consistently located in separate medial and lateral zones. In contrast, no consistent differences in the mediolateral location of label was noted for caudal motor cortex. To summarize, laminae I-III received input only from the somatic sensory cortex, while laminae IV-V received input from both somatic sensory and caudal motor cortex. Lamina VI received input from all cortical fields examined. Laminae VII-IX received input selectively from the rostral motor cortex. For motor cortex, our findings suggest that projections from the two subregions comprise separate descending pathways that could play distinct functional roles in movement control and sensorimotor integration.

Animals↗

Spatial frequency processing in posteromedial lateral suprasylvian cortex does not depend on the projections from the striate-recipient zone of the cat's lateral posterior-pulvinar complex.

It is generally considered that the posteromedial part of the cat's lateral suprasylvian cortex is involved in the analysis of image motion. The main afferents of the posteromedial lateral suprasylvian cortex come from a direct retinogeniculate pathway and indirect retinotectal and retino-geniculo-cortical pathways. Removal of the primary visual cortex does not affect the spatial and temporal processing of suprasylvian cortex cells suggesting that these properties are derived from thalamic input. We have investigated the possibility that the striate-recipient zone of the lateral posterior nucleus-pulvinar complex may be responsible for the spatial (and temporal) frequency processing in posteromedial lateral suprasylvian cortex since these two regions establish strong bidirectional connections and share many visual properties. Experiments were done on anaesthetized normal adult cats. Visual responses in suprasylvian cortex were recorded before, during, and after the deactivation of the lateral part of the lateral posterior nucleus accomplished by the injection of lidocaine or GABA. Results can be summarized as follows. A total of 64 cells was tested. Out of this number, 11 units were affected by the deactivation of the lateral part of lateral posterior nucleus and one cell, by the blockade of pulvinar. For all cells, except one, the effect consisted in a global reduction of the evoked discharge rate suggesting that the thalamo-suprasylvian cortex projections are excitatory in nature. We did not find any significant differences in the optimal spatial frequency, nor in the width of the tuning function, whether the grating was presented at half- or saturation contrast. In addition, there were no significant differences between the low- and high cut-off spatial frequency values computed before and after the deactivation of the lateral posterior nucleus. No specific changes were observed in the contrast sensitivity function of the posteromedial lateral suprasylvian cortex cells. Similar results were observed with respect to the temporal frequency tuning functions. Deactivating the lateral posterior nucleus did not modify the direction selectivity nor the organization of the subregions of the lateral suprasylvian cortex "classical" receptive fields. The absence of strong changes in posteromedial lateral suprasylvian cortex cell response properties following the functional blockade of the lateral posterior nucleus suggests that the projections from this part of the thalamus are not essential to generate the spatial characteristics of most posteromedial lateral suprasylvian cortex receptive fields. These properties may be derived from other thalamic inputs (e.g., medial interlaminar nucleus) and/or from the intrinsic computation of the afferent signals within the lateral suprasylvian cortex. On the other hand, it is possible that the lateral posterior nucleus lateral suprasylvian cortex loop may be involved in other functions such as the analysis of complex motion as suggested by the findings from our and other groups.

Animals↗

Potentials evoked in human and monkey cerebral cortex by stimulation of the median nerve. A review of scalp and intracranial recordings.

Somatosensory evoked potentials (SEPs) are generated in afferent pathways, subcortical structures and various regions of cerebellar and cerebral cortex by stimulation of somatic receptors or electrical stimulation of peripheral nerves. This review summarizes current knowledge of SEPs generated in cerebral cortex by stimulation of the median nerve, the most common form of stimulation for human research and clinical investigations. Major sources of data for the review are intracranial recordings obtained from patients during diagnostic or neurosurgical procedures, and similar recordings in monkeys. Short-latency cortical SEPs in the 20-40 ms latency range consist of P20 and N30, recorded from motor cortex and frontal scalp; P25 and N35, recorded from cortex near the central sulcus and central scalp; and N20 and P30, recorded from somatosensory cortex and parietal scalp. Several lines of evidence including cortical surface and intracerebral recordings, neuromagnetic recordings and lesion studies in humans and monkeys, strongly support the conclusion that these potentials are generated in contralateral somatosensory cortex in areas 3b and 1, in contrast to the conclusion of many previous studies that SEPs recorded from the frontal scalp are generated in motor cortex and other frontal lobe areas. These potentials are primarily mediated by cutaneous afferents of the dorsal column-medial lemniscal system; the contribution of muscle afferents has not been completely resolved but appears to be small. There is currently no evidence that short-latency SEPs are generated in cortex other than primary somatosensory cortex. Recordings from the vicinity of the second somatosensory area, from the supplementary motor and sensory areas and from surface cortex other than sensorimotor cortex have not detected reliable short-latency activity, although some of these regions generate long-latency potentials. Consequently, short-latency SEPs recorded from the scalp are similar to those recorded from the surface of sensorimotor cortex. Old World monkeys such as Macaca mulatta and M. fascicularis provide an excellent model for human short-latency SEPs. All the potentials described above in humans have corresponding monkey analogues, with similar distributions over the cortical surface. The squirrel monkey, a New World species, exhibits the same potentials, but due to the different morphology of sensorimotor cortex, the surface distribution of SEPs is quite different.

Afferent Pathways↗

Phenotypic characterisation of respecified visual cortex subsequent to prenatal enucleation in the monkey: development of acetylcholinesterase and cytochrome oxidase patterns.

Prenatal bilateral enucleation induces cortex, which normally would have become striate cortex, to follow a default developmental pathway and to take on the cytoarchitectonic appearance of extrastriate cortex (default extrastriate cortex, Dehay et al. [1996] J. Comp. Neurol. 367:70-89). We have investigated if this manipulation influences the cortical expression of acetylcholinesterase (AChE) and cytochrome oxidase (CO). Early enucleation (before embryonic day 81; E81) had only minor effects on the distribution of AChE and CO in the striate cortex. In animals that underwent operation, the striate cortex CO blobs were significantly more closely spaced on the operculum compared with the calcarine. After early enucleation, there was a periodic distribution of CO dense patches in default extrastriate cortex. These CO patches had a center-to-center spacing that was considerably smaller than that of CO stripes in normal area V2, but was somewhat larger than that of the CO blobs in striate cortex. Although the CO stripes characteristic of normal area V2 could not be detected, there were some high-frequency CO patches, similar to those found in default extrastriate cortex. Early enucleation caused a failure to form the transient AChE bands running perpendicular to the striate border, which are normally present in the fetus and early neonate. Late enucleation did not alter AChE expression in extrastriate cortex. The relatively minor effects of early enucleation in the reduced striate cortex contrast with the changes in expression of these enzymes in extrastriate cortex, which accompany large shifts in the location of the striate border. This suggests a massive reorganisation of cortical phenotype in extrastriate cortex.

Acetylcholinesterase↗

Direct projection from the dorsal lateral geniculate nucleus to the prestriate cortex in macaque monkeys.

The enzyme horseradish peroxidase (HRP) was separately injected into striate, prestriate, inferotemporal, and parietal cortices in 19 macaque monkeys, and the lateral geniculate nucleus (LGN) was examined for retrograde transport. Labeled LGN cells were identified only in the animals, with HRP injections into the striate and prestriate cortex. Following injections into either of these regions, labeled cells were found in both parvocellular and magnocellular regions of the ipsilateral LGN only, in keeping with the topographic relation of HRP injection sites in the cortex to labeled areas in the LGN. It was also found that (1) labeled LGN cells were less numerous in both laminar and interlaminar zones following HRP injection into the prestriate cortex, whereas following HRP injection into the striate cortex labeled cells were found almost exclusively in the laminae, and localized to a wedge-shaped region; (2) following HRP injection into the prestriate cortex, the mean sizes of the labeled parvocellular and magnocellular cells, estimated in projected diameter, were almost the same, these means being significantly larger than the mean size of labeled parvocellular cells and much smaller than that of labeled magnocellular cells following HRP injection into the striate cortex; (3) the shapes of the labeled LGN cells following HRP injection into the prestriate cortex were ovoid, fusiform, or triangular (or multipolar), whereas those following HRP injection into the striate cortex were uniformly ovoid or round. The above findings following HRP injections into the prestriate cortex in normal monkeys were confirmed by HRP injections into the prestriate cortex of monkeys whose striate cortex had been removed several months prior to the injection; labeled cells were found in confines of areas of retrograde degeneration in the LGN and their labeling pattern was the same as that in intact animals. It was concluded that in macaque monkeys, just as in the cat, a geniculoprestriate projection system exists; it was suggested that there are two parallel system of visual information processing from the LGN to the prestriate cortex, a direct one and in indirect one through the striate cortex.

Animals↗

Projections of the dorsal lateral geniculate and lateral posterior nuclei to visual cortex in the rabbit.

The origin and terminations of thalamic inputs to the striate cortex and the occipital cortex of the rabbit were studied using both anterograde autoradiographic techniques and retrograde transport of horseradish peroxidase (HRP). After injections of [3H]-leucine into the dorsal lateral geniculate nucleus (DLGN) the transport of radiolabeled material was demonstrated in separate loci in both the striate and the occipital cortex. In both these cortical areas, the principal site of geniculocortical termination was in lamina IV with some diminished input spreading into laminae II-III and a light termination in layer I overlying the lamina IV termination. Layer VI of striate cortex received a substantial projection from DLGN while infragranular laminae of occipital cortex received a similar although lighter and more diffuse projection. The lateral posterior nucleus (LPN) was similarly demonstrated to project to both striate and occipital cortices, the projection terminating principally in lamina IV of occipital cortex, lamina V of striate cortex, and layer I over a large, continuous area of the posterior pole of the cortex. Moreover, a projection from LPN to the retrosplenial cortex medial to the striate area was consistently seen. The autoradiographic demonstration of a projection from DLGN and LPN to both striate cortex and occipital cortex was corroborated by the retrograde studies. Following the injection of HRP into either the striate or occipital cortex, columns of retrogradely filled somata were identified in both the DLGN and LPN. The location of the column of labeled neurons within each nucleus varied predictably with the location of the injection in either the striate or the occipital cortex.

Animals↗

Thalamic projections to retrosplenial cortex in the rat.

The topographic relationships between anterior thalamic neurons and their terminal projection fields in the retrosplenial cortex of the rat were characterized by experiments with the fluorescent dye retrograde labeling technique. The results demonstrate that the anterodorsal (DAD) and anteroventral (AV) nuclei project heavily to retrosplenial granular cortex (Rg) and to a lesser extent to retrosplenial agranular cortex (Rag). In contrast, the anteromedial (AM) and lateral dorsal (LD) nuclei project heavily to Rag and more lightly to Rg. Irrespective of terminal field in Rg or Rag, the neuronal cell bodies in AD and AV are organized topographically so that the neurons in the caudal part of each nucleus project to rostral retrosplenial cortex and the neurons in the rostral portion of each nucleus project to the caudal retrosplenial cortex. Further, the ventromedial AD and AV neurons project to rostral retrosplenial cortex, whereas dorsolateral neurons in both nuclei project to caudal retrosplenial cortex. LD neurons display a different topographic organization. The neurons in the medioventral part of LD project primarily to the rostral retrosplenial cortex, and the neurons in lateral LD project to the caudal retrosplenial cortex. This latter projection to the caudal retrosplenial cortex is also contributed to by neurons residing in the mediodorsal part of caudal LD. The neurons in AM that project to the retrosplenial cortex display less segregation than the AV, AD, or LD neurons. In all experiments, a number of neurons in the dorsal ventro-anterolateral nucleus were labeled by retrosplenial injections. The largest number of cells in this nucleus were labeled after Rag injections, and these were topographically organized such that the neurons projecting to the rostral Rag were located immediately deep to the internal medullary lamina, and the neurons projecting to the caudal Rag were more ventrally located. Very few thalamic neurons have axon collaterals to different areas of the retrosplenial cortex as shown by double labeling experiments. Together, these results demonstrate a highly organized thalamic projection to the retrosplenial cortex.

Animals↗

Cortical connections of electrophysiologically and architectonically defined subdivisions of auditory cortex in squirrels.

Multiunit recordings with microelectrodes were used to identify and delimit subdivision of auditory cortex in squirrels. In the same animals, cortical connections of subdivisions of auditory cortex were determined by placing injections of the tracer wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP) into electrophysiologically defined locations. The electrophysiological results and patterns of connections were later related to myeloarchitectonic distinctions in brain sections cut parallel to the surface of the artificially flattened cortex. As previously described (Merzenich et al.: J. Comp. Neurol. 166:387-402, '76), a primary auditory field, A-I, was characterized by (1) neurons narrowly tuned to tone frequency; (2) a tonotopic map with high frequencies, which represented caudal to low frequencies; and (3) dense myelination. A-I was reciprocally connected with a rostral field, R, a parietal ventral somatosensory representation, PV, cortex ventral to A-I, and other nearby regions of cortex of the same hemisphere. Callosal connections of A-I were with A-I, R, and two or more other regions of temporal cortex. The less densely myelinated rostral field, R, also had neurons that were frequency tuned, but the neurons were often less securely driven. R appeared to have a tonotopic organization that roughly mirrored that of A-I. Ipsilateral connections of R included A-I, PV, and cortex ventral and caudal to R. Callosal connections were with R, A-I, PV, and cortex ventral and caudal to R. Callosal connections were with R, A-I, PV, and other locations in temporal cortex. Cortex in caudal PV, ventral to A-I, and ventral to R was responsive to auditory stimuli, but responses to pure tones were weak and inconsistent, and habituation to a repeated stimulus was rapid. The cortex responsive to auditory stimuli included some but not all of the cortex connected with A-I and R. The results lead to the conclusion that auditory cortex of squirrels contains at least two tonotopically organized fields, possibly as many as five or more auditory fields, and at least two auditory-somatosensory fields.

Animals↗

Thalamic projections to the lateral suprasylvian visual area in cats with neonatal or adult visual cortex damage.

Previous transneuronal anterograde tracing studies have shown that the retino-thalamic pathway to the posteromedial lateral suprasylvian (PMLS) visual area of cortex is heavier than normal in adult cats that received neonatal damage to visual cortical areas 17, 18, and 19. In contrast, the strength of this projection does not appear to differ from that in normal animals in cats that experienced visual cortex damage as adults. In the present study, we used retrograde tracing methods to identify the thalamic cells that project to the PMLS cortex in adult cats that had received a lesion of visual cortex during infancy or adulthood. In five kittens, a unilateral visual cortex lesion was made on the day of birth, and horseradish peroxidase (HRP) was injected into the PMLS cortex of both hemispheres when the animals were 10.5 to 13 months old. For comparison, HRP was injected bilaterally into the PMLS cortex of three cats 6.5 to 13.5 months after they received a similar unilateral visual cortex lesion as adults. In cats with a neonatal lesion, retrograde labeling was found in the large neurons that survive in the otherwise degenerated layers A and A1 of the lateral geniculate nucleus (LGN) ipsilateral to the lesion. Retrograde labeling of A-layer neurons was not seen in the undamaged hemisphere of these animals or in either hemisphere of animals that had received a lesion as adults. As in normal adult cats, retrograde labeling also was present in the C layers of the LGN, the medial interlaminar nucleus, the posterior nucleus of Rioch, the lateral posterior nucleus, and the pulvinar nucleus ipsilateral to a neonatal or adult lesion. Quantitative estimates indicate that the number of labeled cells is much larger than normal in the C layers of the LGN ipsilateral to a neonatal visual cortex lesion. Thus the results indicate that the heavier than normal projection from the thalamus to PMLS cortex that exists in adult cats after neonatal visual cortex damage arises, at least in part, from surviving LGN neurons in the A and C layers of the LGN. Although several thalamic nuclei, as well as the C layers of the LGN, continue to project to PMLS cortex after an adult visual cortex lesion, these projections appear not to be affected significantly by the lesion.

Aging↗

Heterogeneity of layer II neurons in human entorhinal cortex.

Abnormalities in the layer II neurons of human entorhinal cortex have been implicated in the pathophysiology of Alzheimer's disease and schizophrenia. The reported abnormalities are not homogeneously distributed throughout the entorhinal cortex, suggesting that layer II of entorhinal cortex may contain different subpopulations of neurons, each with a different susceptibility to pathological mechanisms. In order to investigate the possible heterogeneity of neurons in layer II of human entorhinal cortex, we first identified distinct subdivisions of human entorhinal cortex by adapting the cytoarchitectonic criteria for subdivisions of monkey entorhinal cortex described by Amaral et al. (J Comp Neurol 264:326, 1987). The morphology and regional distribution of distinct subpopulations of human layer II neurons were determined through the use of immunohistochemical techniques. Multipolar, stellate, and modified pyramidal neurons in the characteristic cell clusters or islands of layer II were immunoreactive for nonphosphorylated neurofilament proteins. The intensity of immunoreactivity for the nonphosphorylated neurofilament proteins gradually increased along the rostrocaudal axis of entorhinal cortex and was primarily due to a similar gradient in the density of labeled neurons per island. The calcium-binding protein calbindin D-28K was found in both pyramidal and nonpyramidal neurons in layers II and superficial III. The distribution of calbindin-immunoreactive neurons also depended upon the region of entorhinal cortex. In rostral entorhinal cortex, labeled neurons were scattered throughout the superficial layers, whereas in caudal entorhinal cortex, distinctive patches of small calbindin-immunoreactive neurons were found among the layer II islands. Another calcium-binding protein, parvalbumin, was present in nonpyramidal neurons in layers II and III that were distinct from those containing calbindin. The regional distribution of parvalbumin-positive neurons was very similar to that of the neurofilament immunoreactive neurons; in rostral entorhinal cortex very few parvalbumin-labeled neurons were present but their frequency gradually increased in the caudal direction. In addition, punctate parvalbumin immunoreactivity was frequently encountered in the location of the nonphosphorylated neurofilament protein-positive layer II islands. These findings demonstrate that layer II of human entorhinal cortex contains distinct subpopulations of neurons, that the relative density of each subpopulation differs across cytoarchitectonic regions, and that the patterns of distribution of these subpopulations are in some cases similar and in other cases complementary. This heterogeneity in the organization of layer II of human entorhinal cortex has important implications for the study of some neuropsychiatric disorders.

Adult↗

Cortical connections of subdivisions of inferior temporal cortex in squirrel monkeys.

Patterns of cortical connections and architectonics were used to determine subdivisions of inferior temporal (IT) cortex of squirrel monkeys. Single or multiple injections of the tracers wheat germ agglutinin-horseradish peroxidase, Fast Blue, Diamidino Yellow, Fluoro-Gold, and 3H-amino acids were placed into IT cortex. Most injections were placed in caudal IT cortex in the region previously shown to receive input from the caudal subdivision of the Dorsolateral Area, DLC; additional injections were placed in more rostral IT cortex. The results indicate the presence of two major regions: a caudal region, ITC, and a rostral region, ITR. An intermediate region of cortex along the ITC-ITR border that displays some connections of ITC and some connections of ITR may be another area. ITC contains a more myelinated dorsal area, ITCd, and a larger ventral area, ITCv. Both ITCd and ITCv receive a major projection from DLC; additional input from DLR, MT, and VII; and send strong projections to ITR, the lateral bank of the superior temporal sulcus, and dorsolateral prefrontal cortex. Only ITCd has strong connections with DLR and cortex in the depths of the superior temporal sulcus, and only ITCv has connections with lateral orbital cortex. The overall pattern of connections between ITC and DLC suggests that ITC has a crude topographic organization, with dorsal cortex representing the lower field and ventral cortex representing the upper field. ITR differs from ITC by receiving little if any input from DLC; projecting to inferior temporal polar cortex, the rostral Sylvian fissure, and medial orbital cortex; and having a less distinct layer IV. Comparison of subdivisions of inferior temporal cortex defined in the present study in squirrel monkeys and those reported in other primates suggests that ITC of squirrel monkeys may correspond to area TEO of macaque monkeys.

Animals↗

Effects of neonatal enucleation on the organization of callosal linkages in striate cortex of the rat.

Lewis and Olavarria ([1995] J. Comp. Neurol. 361:119-137) showed that the mediolateral organization of callosal linkages differs markedly between medial and lateral regions of striate cortex in the rat. Thus, callosal fibers originating from medial regions of striate cortex interconnect loci that are mirror-symmetric with respect to the midsagittal plane. In contrast, fibers from lateral regions of striate cortex show a reversed pattern of connections: tracer injections into the 17/18a border produce retrograde cell labeling in regions medial to the contralateral 17/18a border, whereas injections placed somewhat medial to the 17/18a border label cells located at the contralateral 17/18a border. Based on the interpretation that callosal fibers from lateral striate cortex connect retinotopically corresponding loci (Lewis and Olavarria [1995] J. Comp. Neurol. 361:119-137) we propose here that the development of the reversed pattern of connections in lateral portions of striate cortex is guided by activity-dependent cues originating from spontaneously active ganglion cells in temporal retina. In the present study we have attempted to falsify this hypothesis by investigating the effects of neonatal bilateral enucleation on the organization of callosal linkages in striate cortex of the rat. Once enucleated rats reached adulthood, we studied the mediolateral organization of callosal connections by placing small injections of different fluorescent tracers into different loci within medial and lateral striate cortex. The analysis of the distribution of retrogradely labeled callosal cells indicated that connections from lateral portions of striate cortex were no longer organized in a reversed fashion, rather, they resembled the mirror image pattern normally found in the medial callosal region, i.e., injections at the 17/18a border produced labeled cells at the opposite 17/18a border, whereas injections into slightly more medial regions produced labeled cells in the opposite, mirror-symmetric location. In addition, we found that enucleation does not alter the organization of callosal linkages in medial portions of striate cortex. Thus, by showing that enucleation significantly changes the pattern of connections from lateral portions of striate cortex, the present study does not falsify, but rather strengthens the hypothesis that interhemispheric correlated activity driven from the temporal retinal crescent guides the normal development of reversed callosal linkages in lateral portions of rat striate cortex. Furthermore, the present study shows that, in the absence of the eyes, the pattern of callosal linkages in lateral portions of striate cortex resembles the mirror image pattern normally found only in medial striate cortex.

Animals↗

Contribution of striate inputs to the visuospatial functions of parieto-preoccipital cortex in monkeys.

In Experiments 1 and 2, monkeys received 3-stage operations intended to serially disconnect parieto-preoccipital from striate cortex. At each stage (unilateral parieto-preoccipital removal, contralateral striate removal and posterior callosal transection) the monkeys were tested for retention of the landmark task, a visuospatial discrimination sensitive to the effects of bilateral parieto-preoccipital damage. To check the effectiveness of the disconnection, the monkeys were also tested after removal of the remaining parieto-preoccipital cortex. The results demonstrated that corticocortical inputs from striate cortex are crucial for the visuospatial functions of parieto-preoccipital cortex, just as they had been shown earlier to be crucial for the pattern discrimination functions of inferior temporal cortex. Relative to inferior temporal cortex, however, parieto-preoccipital cortex was found to be especially dependent on ipsilateral (as compared with contralateral) striate inputs. In Experiment 3, monkeys received bilateral lesions of either lateral on medial striate cortex and were tested on both a pattern discrimination task, to assess residual inferior temporal function, and the landmark task, to assess residual parieto-preoccipital function. The results indicated that the pattern discrimination functions of inferior temporal cortex are especially dependent on inputs from lateral striate cortex, whereas the visuospatial functions of parieto-preoccipital cortex are equally dependent on inputs from lateral and medial striate cortex. The relatively greater contribution to parieto-preoccipital than to inferior temporal cortex made by ipsilateral and medial striate inputs (representing contralateral and peripheral visual fields, respectively) can also be seen in the receptive field properties of parieto-preoccipital and inferior temporal neurons. The differences in the organization of striate inputs to these two cortical association areas presumably reflect differences in the processing required for spatial vs object vision.

Animals↗

Neurogenesis in the rat primary olfactory cortex.

Neurogenesis in the rat primary olfactory cortex was examined with [3H]thymidine autoradiography. The experimental animals were the offspring of pregnant females given an injection of [3H]thymidine on two consecutive gestation days. Nine groups of embryos were exposed to [3H]thymidine on E13-E14, E14-E15....E21-E22, respectively. On P60, the percentage of labeled cells and the proportion of cells originating during 24 hr periods were quantified at selected anatomical levels of the anterior and posterior piriform cortex, dorsal lateral peduncular cortex, and posterior two-thirds of the ventral agranular insular cortex. Throughout most of the primary olfactory cortex, deep cells are generated earlier than superficial cells: the 'inside-out' pattern. Neurons in the anterior (prepiriform) cortex are located lateral to the caudal anterior olfactory nucleus and olfactory tubercle, and are generated mainly between E14 and E18 in a caudal (older) to rostral (younger) neurogenetic gradient. Neurons in the posterior (periamygdaloid) cortex are located lateral to the caudal olfactory tubercle and amygdala, and are generated mainly between E14 and E17 simultaneously along the rostrocaudal plane. Superficial cells in the piriform cortex have some additional neurogenetic gradients; ventromedial cells forming transition zones with either the olfactory tubercle or amygdala originate earlier than cells located dorsally and laterally. In the posterior piriform cortex, younger neurons are located at middle dorsoventral levels while older neurons lie above and below. Neurons in the dorsolateral peduncular cortex originate between E14 and E20 in a caudal to rostral gradient of neurogenesis; caudal parts also have a lateral to medial neurogenetic gradient. The most lateral part of the dorsolateral peduncular cortex is unique and does not have the typical 'inside-out' cortical neurogenetic gradient. Neurons in the ventral agranular insular cortex (area 13) originate mainly between E15 and E17 in combined caudal to rostral and ventral to dorsal neurogenetic gradients. The neurogenetic gradients in the primary olfactory cortex, along with patterns of neurogenesis throughout the olfactory projection field are related to the termination patterns of afferents from the main olfactory bulb.

Animals↗

To-and-fro optical voltage signal propagation between the insular gustatory and parietal oral somatosensory areas in rat cortex slices.

Taste perception depends not only on special taste information processed in the insular cortex, but also on oral somesthetic processing in the parietal cortex. Many insular cortex neurons show multimodal responsiveness. Such multimodality may be enabled by signal exchange between these two cortices. By using the protocol that we have developed, a synchronized population oscillation of synaptic potentials was induced in the parietal cortex by stimulation to the insular cortex in rat neocortex slices. The spatiotemporal pattern of propagation of this oscillation was studied by recording voltage-sensitive optical signals and field potentials. The first wavelet of the oscillation was propagated from the insular stimulation site to the parietal cortex. However, the second and later wavelets propagated back from the parietal cortex to the insular cortex. The oscillation was detected in the insular cortex as well, but was actually generated in the parietal cortex. Thus, the initial peak of optical signal, sent from the insular to parietal cortex, served to generate oscillatory responses in the parietal cortex, which propagated back to the insular cortex wave-by-wave. We propose that this to-and-fro propagation may be an artificially exaggerated demonstration of an intrinsic mechanism relevant to signal exchange between the parietal and insular cortices.

Animals↗

Transient induction of neuronal nitric oxide synthase in neurons of rat cerebral cortex after status epilepticus.

The change in neuronal nitric oxide synthase (nNOS) expression after status epilepticus induced by kainate was examined in the rat cerebral cortex. Expressional change was assessed using nNOS immunohistochemistry and reverse transcription-polymerase chain reaction for nNOS mRNA. Constitutive nNOS-positive neurons was observed in the cerebral cortex of the control group. At 1 and 3 days after status epilepticus, nNOS-positive neurons were present in a deep layer of various cortical regions such as primary motor cortex, secondary motor cortex, primary somatosensory cortex, secondary somatosensory cortex, parietal association cortex, insular cortex, ectorhinal cortex, temporal association cortex, auditory cortex and visual cortex. The level of nNOS mRNA increased at 1, 3, 6 and 12 days after status epilepticus compared to controls. This report provides the first morphological evidence that nNOS are induced in neurons of the cerebral cortex following seizure.

Animals↗

Age-related change in the neuropeptide Y and NADPH-diaphorase-positive neurons in the cerebral cortex and striatum of aged rats.

Age-related changes of neuropeptide Y (NPY) and nicotinamide adenine dinucleotide phosphate-diaphorase (NADPH-d) were examined in the rat cerebral cortex and striatum by immunohistochemical and histochemical methods. Double labeling for NPY and NADPH-d showed that about 30-70% of NPY-immunoreactive (NPY-IR) neurons in the cerebral cortex of the control (4-month-old) rats contained NADPH-d and that 50-75% in the aged (24-month-old) rats. The aged rats showed a significant increase in percentage of colocalization of NPY and NADPH-d in comparison with the control rats in the temporal cortex, occipital cortex, cingulate cortex, insular cortex, retrosplenial cortex and caudatoputamen. However, colocalization percentage between control and aged rats in the frontal cortex, parietal cortex, perirhinal cortex, entorhinal cortex and nucleus accumbens were practically identical. In the aged group, the number of NPY-IR/NADPH-d-positive neurons was not significantly decreased in the cerebral cortex and striatum compared to the control group. However, the number of NPY-IR/NADPH-d-negative neurons was significantly decreased in all cerebral cortical areas and caudatoputamen in the aged group except in the nucleus accumbens. Major loss of NPY-IR/NADPH-d-negative neurons in the aged group were observed in the neurons of layer II/III and V/VI. These results demonstrate that the NADPH-d containing NPY-IR neurons are less influenced by aging than the control group in the cerebral cortex and striatum of rats.

Aging↗