Induction of c-fos protein by patterned visual stimulation in central visual pathways of the rat

Localized patterned visual stimulation was used in rats to investigate the feasibility of stimulus-dependent induction of the immediate early gene c-fos in neurons of cortical and subcortical visual centers of this mammal. Moving and stationary visual patterns, consisting of gratings and arrays of dark dots, induced Fos-like immunoreactivity in populations of neurons in retinotopically corresponding stimulated regions of the dorsal and ventral lateral geniculate nucleus (dLGN, vLGN), stratum griseum superficiale of the superior colliculus, nucleus of the optic tract, and primary (striate) visual cortex. Only moving stimuli induced Fos-like immunoreactive (FLI) neurons in extrastriate visual areas, particularly in the anterolateral (AL) visual area. This suggests that area AL is equivalent to the motion sensitive areas MT and PMLS of the monkey and cat. Stimulus-induced FLI neurons in the striate cortex were predominantly distributed in layers 4 and 6, while few labeled neurons were present in layers 2–3, and almost none in layer 5. The laminar distribution of stimulus-induced FLI cells in the extrastriate cortical area AL was similar to that of the striate cortex, with the exception that more FLI cells were present in layer 5. Statistical comparison of somata size of the stimulus-induced FLI neurons in dLGN with that of Cresyl violet stained neurons in the same sections revealed that the population of geniculate FLI neurons is composed of relay cells and interneurons.     

Monoclonal antibody to neurofilament protein (SMI-32) labels a subpopulation of pyramidal neurons in the human and monkey neocortex

A monoclonal antibody that recognizes a nonphosphorylated epitope on the 168 kDa and 200 kDa subunits of neurofilament proteins has been used in an immunohistochemical study of cynomolgus monkey (Macaca fascicularis) and human neocortex. This antibody, SMI-32, primarily labels the cell body and dendrites of a subset of pyramidal neurons in both species. A greater proportion of neocortical pyramidal neurons were SMI-32 immunoreactive (ir) in the human than in the monkey. SMI-32-ir neurons exhibited consistent differences in the intensity of their immunoreactivity that correlated with cell size. The cellular specificity of SMI-32 immunoreactivity suggests that a subpopulation of neurons can be distinguished on the basis of differences in the molecular characteristics of basic cytoskeletal elements such as neurofilament proteins. The size, density, and laminar distribution of SMI-32-ir neurons differed substantially across neocortical areas within each species and between species. Differences across cortical areas were particularly striking in the monkey. For example, the anterior parainsular cortex had a substantial population of large SMI-32-ir neurons in layer V and a near absence of any immunoreactive neurons in the supragranular layers. This contrasted with the cortical area located more laterally on the superior temporal gyrus, where layers III and V contained substantial populations of large SMI-32-ir neurons. Both areas differed significantly from the posterior inferior temporal gyrus, which was distinguished by a bimodal distribution of large SMI-32-ir neurons in layer III. Differences across human areas were less obvious because of the increase in the number of SMI-32-ir neurons. Perhaps the most notable differences across human areas resulted from shifts in the density of the larger SMI-32-ir neurons in deep layer III. A comparison between the species revealed that isocortical areas exhibited greater differences in their representation of SMI-32-ir neurons than primary sensory or transitional cortical areas. A comparison of distribution patterns of SMI-32-ir neurons across monkey cortical areas and data available on the laminar organization of cortical efferent neurons suggests that a common anatomic characteristic of this chemically identified subpopulation of neurons is that they have a distant axonal projection. Such correlations of cell biological characteristics with specific elements of cortical circuitry will further our understanding of the molecular and cellular properties that are critically linked to a given neuron's role in cortical structure and function.     

Capacity of the retinogeniculate pathway to reorganize following ablation of visual cortical areas in developing and mature cats

The purpose of the present study was to determine the pattern and density of retinal projections to the dorsal lateral geniculate nucleus (dLGN) following ablation of visual cortical areas in developing cats of different postnatal ages and in mature cats. The terminations of retinal projections to the dLGN were evaluated following the injection of tritiated amino acids into one eye. Regardless of age, a visual cortical ablation of areas 17 and 18 induces massive death of neurons within the regions of the dLGN that are linked topographically to the cortical areas removed. However, the pattern of retinal projections to these degenerated regions of the dLGN differs depending upon whether the cortical lesion is incurred early in postnatal life or in adulthood. Following ablation on the day of birth (P1), virtually all surviving cells were found in the C-complex of dLGN with only a token number in the A-laminae. Correspondingly, retinal projections were maintained to the C-complex of the nucleus and were barely detectable in the degenerated A-laminae. However, in cats in which areas 17 and 18 had been removed in adulthood (≥ 6 months of age) retinal projections were maintained to the A-laminae even though nearly all neurons in those laminae had degenerated. Moreover, a subgroup of animals that incurred area 17 and 18 ablations at P1 showed that the modification of retinal projections to the A-laminae occurs within the first postnatal month, and an additional subgroup showed that retinal projections become increasingly resistant to the degenerative events in the dLGN that follow ablation of areas 17 and 18 at progressively older ages during the first postnatal month. Furthermore, retinal inputs also respond, in an age-dependent way, to degeneration of neurous in the C-complex induced by extension of the cortical ablation to include extrastriate visua areas. © 1993 Wiley-Liss, Inc.     

Thalamo-cortical organization of the visual system in the cat

Thalamo-cortical relationships in the visual system of the cat were studied by the method of retrograde degeneration. Localized lesions limited to area 17 result in degeneration only in the dorsolateral geniculate body; cell changes are marked in 3 laminae (A, A₁, B), mild in nucleus interlaminaris centralis and minimal in nucleus interlaminaris medialis. Lesions limited to areas 18 and 19 are followed by marked degeneration in the medial interlaminar nucleus, mild in the other laminae; in addition, the lateral part of the posterior thalamic nucleus (ventral or inferior pulvinar) is also atrophied. Following large striate lesions which marginally involved areas 18 and 19, there is also mild, localozed degeneration in the anteroventral and reticular thalamic nuclei. Whin cortical lesions are limited to the convexity of the suprasylvian gyri, degeneration is present in the lateral aspect of laminae A, A₁, B and nucleus interlaminaris centralis and in the medial part of the posterior nucleus, in addition to lateral dorsal, lateral posterior and pulvinar nuclei. Lesions in the ectosylvian gyri result in slight but definite degeneration in the lateral part of lamina A of the dorsal lateral geniculate, but nothing in the posterior nucleus. The geniculate projections to areas 17, 18 and 19, to the suprasylvian and ectosylvian gyri all show a rostrocaudal organization. The geniculostriate projection is also topographically organized in a mediolateral manner. Thus, the geniculocortical projection in the cat is not striate specific but spreads over the occipito-temporal cortex at least as far as the acoustic areas of the ectosylvian gyri. In this species the dorsal lateral geniculate body is not a unitary structure but is a complex of nuclei, all of which receive retinal fibers, and the cortical projections of which overlap those of the posterior, lateral dorsal, lateral posterior, pulvinar, medial geniculate, reticular and anterior thalamic nuclei.     

The serotoninergic fibers in the dorsal lateral geniculate nucleus of the cat: distribution and synaptic connections demonstrated with immunocytochemistry

The distribution, morphology, and synaptic contacts of serotoninergic fibers were studied with immunocytochemical methods in the lateral geniculate complex of the cat. The serotonin-immunoreactive fibers are diffusely distributed throughout the main laminae of the dorsal lateral geniculate nucleus (dLGN) and the perigeniculate nucleus (PGN) and reach a particular density in the ventral lateral geniculate nucleus (vLGN). The labeled fibers are in most cases very thin and sometimes varicose. There is no obvious order in their distribution pattern except that they sometimes partially encircle the unlabeled cell bodies of the dLGN. The synaptic connections of the serotoninergic fibers were investigated mainly in the A laminae of the dLGN. Few synaptic complexes were found, most of them with asymmetric morphology. The postsynaptic elements were small dendritic profiles. Perisomatic serotoninergic fibers were seen, but no convincing synaptic contacts were found between labeled fibers and cell somata. In the dLGN, serotoninergic profiles were almost exclusively confined to the extraglomerular neuropile. In the PGN serotoninergic fibers also contacted dendritic profiles and formed asymmetrical synapses, but as in the geniculate, synaptic specializations were very rare.     

Correlations between thalamic projections to different areas of the parietal association cortex of the cat

Projections of thalamic neurons to parietal association cortex of cat were examined by means of the retrograde axonal transport of fluorescent dyes (primuline and fast blue). It has been demonstrated that a dorsal part of the pulvinar (PL) and a dorsal part of the caudal area of the lateral posterior nucleus (LP) projected mostly to the middle suprasylvian gyrus (MSSG), while a ventral part of PL and a ventral part of the rostral area of LP--to the rostral suprasylvian gyrus (RSSG). Double labelled neurons were found in PL, LP, suprageniculatus, anterior ventral, ventral lateral as well as in the central lateral, paracentral and central medial nuclei after injections of two different dyes into MSSG and RSSG. Topic organization of sources of cortical projections from the PL--LP complex can probably provide a high level of discrimination of visual signals by single cortical neurons. At the same time during integration of information of a different subcortical origin RSSG and MSSG act, probably, in concord to a considerable extent, that suggests insufficient differentiation of RSSG and MSSG corresponding approximately to cortical areas 5 and 7 of cat.     

Differential expression of neurofilament protein in the visual system of the vervet monkey

It has been previously reported that the monoclonal antibody SMI-32 reveals a characteristic pattern of immunostaining which may be used to delineate various cortical modules in the monkey visual system. We wished to examine staining patterns with this antibody at both the lateral geniculate nucleus (LGN) and cortical levels with regard to magno- and parvocellular processing schemes in the vervet monkey. Using standard immunohistochemical procedures, we have found that the M-layers of the LGN were intensely stained in comparison to P-layers and that there were regional variations in staining within the visual cortex that reflected this input. The transition between areas Vl and V2 was especially prominent due to differences in the laminar staining profiles. Another striking result was found within the superior temporal sulcus where heavy SMI-32 immunostaining confined to the floor of the sulcus coincided with a similar zone of intense myelin staining. We have also found a number of other areas within the intraparietal and lateral sulci that show foci of heavy SMI-32 staining. As with Cat-301 immunostaining, the regional variabilities that are observed with SMI-32 in the visual cortex reflect molecular distinctions that may provide further criteria for functional segmentation.     

Cyto- and chemoarchitecture of the cerebral cortex of the Australian echidna (Tachyglossus aculeatus). I. Areal organization

We have examined the topography of the cerebral cortex of the Australian echidna (Tachyglossus aculeatus), using Nissl and myelin staining, immunoreactivity for parvalbumin, calbindin, and nonphosphorylated neurofilament protein (SMI-32 antibody), and histochemistry for acetylcholinesterase (AChE) and NADPH diaphorase. Myelinated fibers terminating in layer IV of the cortex were abundant in the primary sensory cortical areas (areas S1, R, and PV of somatosensory cortex; primary visual cortex) as well as the frontal cortex. Parvalbumin immunoreactivity was particularly intense in the neuropil and somata of somatosensory regions (S1, R, and PV areas) but was poor in motor cortex. Immunoreactivity with the SMI-32 antibody was largely confined to a single sublayer of layer V pyramidal neurons in discrete subregions of the somatosensory, visual, and auditory cortices, as well as a large field in the frontal cortex (Fr1). Surprisingly, SMI-32 neurons were absent from the motor cortex. In AChE preparations, S1, R, V1, and A regions displayed intense reactivity in supragranular layers. Our findings indicate that there is substantial regional differentiation in the expanded frontal cortex of this monotreme. Although we agree with many of the boundaries identified by previous authors in this unusual mammal (Abbie [1940] J. Comp. Neurol. 72:429-467), we present an updated nomenclature for cortical areas that more accurately reflects findings from functional and chemoarchitectural studies.     

Anatomical organization of the primary visual cortex (area 17) of the cat. A comparison with area 17 of the macaque monkey.

Golgi and axonal transport techniques have been used to examine the organization of neurons within primary visual cortex, area 17, of the cat. This organization has been compared to that of the primate cortical area 17 as described in previous studies and it is discussed in relationship to the distribution of afferents from the dorsal lateral geniculate nucleus (dLGN). The visual cortex of the cat and monkey show strong similarities in the laminar positions of neurons projecting extrinsically and also in the restriction of spiny stellate neurons to a central lamina (lamina 4) receiving input from the dLGN. However, lamina 4B in the monkey, which contains spiny stellate neurons but does not receive direct input from the dLGN, has no direct counterpart in cat area 17. Axon projections of spiny stellate neurons in the other divisions of lamina 4 differ in cat and monkey: the small, closely packed neurons in the lowermost division of lamina 4 (4B in the cat, 4Cbeta in the monkey) project chiefly within lamina 4 in the cat whereas in the monkey they have a strong projection to lamina 3. In the cat, spiny stellate neurons of lamina 4A project upon lamina 3 whereas in the monkey those in the apparently equivalent zone, 4Calpha, project upon lamina 4B. Most non-spiny stellate neurons examined have precisely organized interlaminar axonal projections which differ from the axon trajectories of neighboring spiny neurons.     

The Topographic Organization and Axis of Projection within the Visual Sector of the Rabbit's Thalamic Reticular Nucleus

The organization of the visual field representation within the thalamic reticular nucleus (TRN) of the rabbit was studied. Focal injections of horseradish peroxidase (HRP) and/or [3H]proline were made into visuocortical areas V1 and V2 and the dorsal lateral geniculate nucleus (dLGN). The resultant labelling in the thalamus was analysed. A single injection in V1 or V2 results in a single zone of terminal label within the TRN that is restricted to the dorsocaudal part of the sheet-like nucleus. In comparisons of the zones of label following injections at two different cortical sites in V1, a medial to lateral shift in label across the thickness of the TRN sheet is accompanied by a medial to lateral shift in label in the dLGN; a dorsal to ventral shift in label within the plane of the TRN sheet is accompanied by a dorsal to ventral shift in label in the dLGN. Thus, like the dLGN the TRN receives a precise topographic projection from V1. In reconstructions from horizontal sections the zones of label within the TRN resemble 'slabs', which lie within the plane of the nucleus parallel to its borders. Thus, the slabs of visuocortical terminals and reticular dendrites are similarly oriented. As revealed by the orientation of the slabs, the lines of projection representing points in visual space are represented by the oblique rostrocaudal dimension of the TRN. Injections restricted to V1 produce terminal labelling that is confined to the outer two-thirds of the TRN across its thickness, whilst those involving V2 result in terminal labelling within the inner one-third of the nucleus. Thus, the adjacent cortical areas V1 and V2 project in a continuous fashion across the mediolateral dimension of the TRN. The organization of the map within the TRN, which was revealed by visuocortical injections, was confirmed by the pattern of retrograde labelling within the nucleus following geniculate injections of HRP. On the basis of these findings and those in other mammalian species, two major conclusions can be reached that alter our view of the TRN. First, rather than mapping onto the whole nucleus in a continuous fashion, the cortical projection to the TRN has significant discontinuities. Second, rather than integrating efferents from widespread cortical areas, the reticular dendrites are related to focal areas of cortex.     

Chemoarchitectonics and corticocortical terminations within the superior temporal sulcus of the rhesus monkey: Evidence for subdivisions of superior temporal polysensory cortex

Cortex of the upper bank of the superior temporal sulcus (STS) in macaque monkeys, termed the superior temporal polysensory (STP) region, corresponds largely to architectonic area TPO and is connectionally distinct from adjacent visual areas. To investigate whether or not the STP region contains separate subdivisions, immunostaining for parvalbumin and neurofilament protein (using the SMI-32 antibody) was compared with patterns of corticocortical terminations in the STS. Chemoarchitectonic results provided evidence for three caudal-to rostral subdivisions: TPOc, TPOi, and TPOr. Area TPOc was characterized by patchy staining for parvalbumin and SMI-32 in cortical layers IV/III and III, respectively. Area TPOi had more uniform chemoarchitectonic staining, whereas area TPOr had a thicker layer IV than TPOi. The connectional results showed prefrontal cortex in the location of the frontal eye fields (area8) and dorsal area 46 projected in a columnar pattern to all cortical layers of area TPOc, to layer IV of TPOi, and in a columanr fashion, with a moderate increase in density in layer IV, to TPOr. In TPOc, columns of frontal connections showed a peridicity similar to that of the SMI-32 staining. The caudal inferior parietal lobule (area 7a) and superior temporal gyrus projected to each subdivision of area TPO, displaying either panlaminar or fourth-layer terminations. In addition to STP cortex, parvalbumin and SMI-32 immunostaining allowed identification of caudal visual areas of the STS, including MT, MST, FST, and V4t. These areas received first and sixth-layer projections from prefrontal cortex and area 7a. © 1995 Wiley-Liss, Inc.     

Investigation of cytoskeleton proteins in neurons of the cat lateral geniculate nucleus

The gross structure of neurons is supported by proteins that compose the cytoskeleton. Neurofilaments are intermediate cytoskeletal proteins that contribute to neuron structure and function, and three neurofilament subunits different in their molecular mass assemble to form heteropolymers that produce a structure-providing intracellular scaffold. The light neurofilament subunit is obligatory and can assemble with either the medium or heavy subunit, indicating some degree of independence between subunits. The presence of the heavy subunit has been shown to be associated with mature cells and is linked to large neurons in the cerebral cortex and thalamus. Spectrin is a membrane-associated actin-binding protein that, like neurofilament, has been linked to neuron shape. In this study of the cat dorsal lateral geniculate nucleus (dLGN) we examined whether labeling for neurofilament subunits and spectrin is linked to neuron size. We found that about one-third of neurons contained a visible amount of labeling for each neurofilament subunit, and the bulk of these labeled cells were large in comparison to the general population of neurons. The distribution of neuron sizes was not different between neurofilament subunits, indicating that neurofilament subunit content is not determined by neuron size. Spectrin labeling was evident in most dLGN neurons, and was not related to the size of neurons. That reactivity for neurofilament was predominant in large cells led us to directly examine the relationship between neurofilament and interneurons. The large majority of neurofilament-positive neurons did not contain GABA, indicating that neurofilament is predominant in projection cells and not in interneurons.     

Connections of the multiple visual cortical areas with the lateral posterior-pulvinar complex and adjacent thalamic nuclei in the cat

The present report describes the patterns of cat thalamocortical interconnections for each of the 13 retinotopically ordered visual areas and additional visual areas for which no retinotopy has yet emerged. Small injections (75 nl) of a mixture of horseradish peroxidase and [3H]leucine were made through a recording pipette at cortical injection sites identified by retinotopic mapping. The patterns of thalamic label show that the lateral posterior-pulvinar complex of the cat is divided into three distinct functional zones, each of which contains a representation of the visual hemifield and shows unique afferent and efferent connectivity patterns. The pulvinar nucleus projects to areas 19, 20a, 20b, 21a, 21b, 5, 7, the splenial visual area, and the cingulate gyrus. The lateral division of the lateral posterior nucleus projects to areas 17, 18, 19, 20a, 20b, 21a, 21b, and the anterior medial (AMLS), posterior medial (PMLS), and ventral (VLS) lateral suprasylvian areas. The medial division of the lateral posterior nucleus projects to areas AMLS, PMLS, VLS, and the anterior lateral (ALLS), posterior lateral (PLLS), dorsal (DLS) lateral suprasylvian areas, and the posterior suprasylvian areas. In addition, many of these visual areas are also interconnected with subdivisions of the dorsal lateral geniculate nucleus (LGd). Every retinotopically ordered cortical area (except ALLS and AMLS) is reciprocally interconnected with the parvocellular C layers of the LGd. The medial intralaminar nucleus of the LGd projects to areas 17, 18, 19, AMLS, and PMLS. Finally, each cortical area (except area 17) receives a projection from thalamic intralaminar nuclei. These results help to define the pathways by which visual information gains access to the vast system of extrastriate cortex in the cat.     

Neurochemical gradients along monkey sensory cortical pathways: calbindin-immunoreactive pyramidal neurons in layers II and III

We examined the distribution of neurons containing immunoreactivity for three calcium-binding proteins, calbindin, parvalbumin and calretinin, as well as nonphosphorylated neurofilament protein, in cortical areas along the ventral and dorsal cortical visual pathways, and in ventrally-directed somatosensory and auditory cortical pathways. Calbindin-immunoreactive pyramidal neurons showed the most prominent regional differences. They were largely restricted to layers II and III and their number monotonically increased from the primary sensory areas to the anteroventral areas along the ventral visual pathway and along the ventrally-directed somatosensory and auditory pathways. The number of calbindin-immunoreactive pyramidal neurons in layers II and III also increased along the dorsal visual pathway, but the number in the last recognized stage of the dorsal visual pathway (area 7a) was significantly smaller than that at the corresponding stage in the ventral visual pathway (TE). The number of calbindin-immunoreactive pyramidal neurons was highest in layers II and III of areas 35/36, TG, and TF/TH, which represent terminal cortical regions of the pathways. These results show neurochemical differences between cortical areas located at early and late stages along serial corticocortical pathways, as well as confirming differences between pyramidal neurons in the supragranular and infragranular layers.     

Organization of cortical and subcortical projections to medial prefrontal cortex in the cat

We have analyzed the cortical and subcortical afferent connections of the medial prefrontal cortex (MPF) in the cat with the specific aim of characterizing subregional variations of afferent connectivity. Thirteen tracer deposits were placed at restricted loci within a cortical district extending from the proreal to the subgenual gyrus. The distribution throughout the forebrain of retrogradely labeled neurons was then analyzed. Within the thalamus, retrogradely labeled neurons were most numerous in the mediodorsal nucleus and in the ventral complex. The projection from each region exhibited continuous topography such that more medial thalamic neurons were labeled by tracer from more ventral and posterior cortical deposits. Marked retrograde labeling without any sign of topographic order occurred in a narrow medioventral sector of the lateroposterior nucleus. Several additional thalamic nuclei contained small numbers of labeled neurons. In a subset of nuclei closely affiliated with the limbic system (the parataenial, paraventricular, reuniens, and basal ventromedial nuclei), retrograde labeling occurred exclusively after deposits at extremely ventral and posterior cortical sites. Within the amygdala, retrogradely labeled neurons occupied the anterior basomedial nucleus, the posterior basolateral nucleus, and a narrow strip of the lateral nucleus immediately adjoining the basolateral nucleus. The number of labeled neurons was greater after more ventral deposits. Very ventral deposits resulted in extensive labeling of the cortical amygdala. Within the cerebral cortex, the distribution of labeled neurons depended on the location of the tracer deposit. Comparatively dorsal deposits produced prominent retrograde transport to the anterior and posterior cingulate areas, to the agranular insula, and to lateral prefrontal cortex. Comparatively ventral deposits gave rise to prominent labeling of the hippocampal subiculum, various parahippocampal areas, and prepiriform cortex. On the basis of afferent connections, it is possible to divide the cat's medial prefrontal cortex into an infralimbic component, MPFil, marked by strong afferents from prepiriform cortex and the cortical amygdala, and a dorsal component, MPFd, without afferents from these structures. Further, within MPFd, it is possible to define an axis, running from ventral and posterior to dorsal and anterior levels, along which limbic afferents gradually become weaker and projections from cortical association areas gradually become stronger.     

Thalamic inputs to visual areas 1 and 2 in the rabbit

Thalamic projections to two cortical representations of the visual field, visual areas 1 and 2 (V1, V2), in the rabbit were studied by using the retrograde transport of horseradish peroxidase (HRP). Physiological guidance was employed to inject small amounts of HRP into topographically defined regions of V1 or V2. Injections restricted to V1 revealed a dense projection from the dorsal lateral geniculate nucleus as well as projections from the pulvinar, the posterior thalamic nucleus, and the ventral lateral nucleus. Injections restricted to V2 revealed projections from the pulvinar, the ventral lateral nucleus, and the posterior thalamic nucleus, but only a slight projection from the dorsal lateral geniculate nucleus. V2, but not V1, receives an input from neurons within the fiber plexus between the dorsal lateral geniculate nucleus and the pulvinar. Finally, the neurons in the lateral geniculate nucleus that project to V2 have larger somata on average than those that project to V1 (means = 18.25 micron vs. 14.04 micron, P less than .001).     

Organization of subcortical pathways for sensory projections to the limbic cortex. I. Subcortical projections to the medial limbic cortex in the rat

Subcortical afferent projections to the medial limbic cortex were examined in the rat by the use of retrograde axonal transport of horseradish peroxidase. Small iontophoretic injections of horseradish peroxidase were placed at various locations within the dorsal and ventral cingulate areas, the dorsal agranular and ventral granular divisions of the retrosplenial cortex and the presubiculum. Somata of afferent neurons in the thalamus and basal forebrain were identified by retrograde labeling. Each of the anterior thalamic nuclei was found to project to several limbic cortical areas, although not with equal density. The anterior dorsal nucleus projects primarily to the presubiculum and ventral retrosplenial cortex; the anterior ventral nucleus projects to the retrosplenial cortex and the presubiculum with apparently similar densities; and the anterior medial nucleus projects primarily to the cingulate areas. The projections from the lateral dorsal nucleus to these limbic cortical areas are organized in a loose topographic fashion. The projection to the presubiculum originates in the most dorsal portion of the lateral dorsal nucleus. The projection to the ventral retrosplenial cortex originates in rostral and medial portions of the nucleus, whereas afferents to the dorsal retrosplenial cortex originate in caudal portions of the lateral dorsal nucleus. The projection to the cingulate originates in the ventral portion of the lateral dorsal nucleus. Other projections from the thalamus originate in the intralaminar and midline nuclei, including the central lateral, central dorsal, central medial, paracentral, reuniens, and paraventricular nuclei, and the ventral medial and ventral anterior nuclei. In addition, projections to the medial limbic cortex from the basal forebrain originate in cells of the nucleus of the diagonal band. Projections to the presubiculum also originate in the medial septum. These results are discussed in regard to convergence of sensory and nonsensory information projecting to the limbic cortex and the types of visual and other sensory information that may be relayed to the limbic cortex by these projections.     

Investigation of cortical reorganization in area 17 and nine extrastriate visual areas through the detection of changes in immediate early gene expression as induced by retinal lesions

The effect of binocular central retinal lesions on the expression of the immediate early genes c-fos and zif268 in the dorsal lateral geniculate nucleus (dLGN) and the visual cortex of adult cats was investigated by in situ hybridization and immunocytochemistry. In the deafferented region of the dLGN, the c-fos mRNA level was decreased within 3 days. The dimensions of the geniculate region showing decreased amounts of c-fos mRNA matched the predictions based on the lesion size and the retinotopic maps of Sanderson ([1971] J. Comp. Neurol. 143:101-118). We did not detect zif268 mRNA in the dLGN. At the cortical level, both c-fos and zif268 mRNA expression decreased in the sensory-deprived region of area 17. In addition, the portions of areas 18, 19, 21a, 21b, and 7, as well as the posterior medial lateral suprasylvian area, the posterior lateral lateral suprasylvian area, the ventral lateral suprasylvian area, and the dorsal lateral suprasylvian area corresponding to the retinal lesions also displayed decreased c-fos and zif268 mRNA levels. Immunocytochemistry revealed similar changes for Zif268 and Fos protein. Three days post lesion, the dimensions of the lesion-affected cortical loci exceeded the predictions in relation to the size of the retinal lesions and the available retinotopic maps. Longer postlesion survival times clearly resulted in a time-dependent restoration of immediate early gene expression from the border to the center of the lesion-affected cortical portions. Our findings represent a new approach for investigating the capacity of adult sensory systems to undergo plastic changes following sensory deprivation and for defining the topographic nature of sensory subcortical and cortical structures.     

Immunohistochemical parcellation of the ferret (Mustela putorius) visual cortex reveals substantial homology with the cat (Felis catus)

Electrophysiological mapping of the adult ferret visual cortex has until now determined the existence of 12 retinotopically distinct areas; however, in the cat, another member of the Carnivora, 20 distinct visual areas have been identified by using retinotopic mapping and immunolabeling. In the present study, the immunohistochemical approach to demarcate the areal boundaries of the adult ferret visual cortex was applied in order to overcome the difficulties in accessing the sulcal surfaces of a small, gyrencephalic brain. Nonphosphorylated neurofilament (NNF) expression profiles were compared with another classical immunostain of cortical nuclei, Cat‐301 chondroitin sulfate proteoglycan (CSPG). Together, these two markers reliably demarcated the borders of the 12 previously defined areas and revealed further arealization beyond those borders to a total of 19 areas: 21a and 21b; the anterolateral, posterolateral, dorsal, and ventral lateral suprasylvian areas (ALLS, PLLS, DLS, and VLS, respectively); and the splenial and cingulate visual areas (SVA and CVA). NNF expression profile and location of the newly defined areas correlate with previously defined areas in the cat. Moreover, NNF and Cat‐301 together revealed discrete expression domains in the posteroparietal (PP) cortex, demarcating four subdivisions in the caudal lateral and medial domains (PPcL and PPcM) and rostral lateral and medial domains (PPrL and PPrM), where only two retinotopic maps have been previously identified (PPc and PPr). Taken together, these studies suggest that NNF and Cat‐301 can illustrate the homology between cortical areas in different species and draw out the principles that have driven evolution of the visual cortex. J. Comp. Neurol. 518:4439–4462, 2010. © 2010 Wiley‐Liss, Inc.     

Organization of the Visual Reticular Thalamic Nucleus of the Rat

The visual sector of the reticular thalamic nucleus has come under some intense scrutiny over recent years, principally because of the key role that the nucleus plays in the processing of visual information. Despite this scrutiny, we know very little of how the connections between the reticular nucleus and the different areas of visual cortex and the different visual dorsal thalamic nuclei are organized. This study examines the patterns of reticular connections with the visual cortex and the dorsal thalamus in the rat, a species where the visual pathways have been well documented. Biotinylated dextran, an anterograde and retrograde tracer, was injected into different visual cortical areas [17; rostral 18a: presumed area AL (anterolateral); caudal 18a: presumed area LM (lateromedial); rostral 18b: presumed area AM (anteromedial); caudal 18b: presumed area PM (posteromedial)] and into the different visual dorsal thalamic nuclei (posterior thalamic, lateral posterior, lateral geniculate nuclei), and the patterns of anterograde and retrograde labelling in the reticular nucleus were examined. From the cortical injections, we find that the visual sector of the reticular nucleus is divided into subsectors that each receive an input from a distinct visual cortical area, with little or no overlap. Further, the resulting pattern of cortical terminations in the reticular nucleus reflects largely the patterns of termination in the dorsal thalamus. That is, each cortical area projects to a largely distinct subsector of the reticular nucleus, as it does to a largely distinct dorsal thalamic nucleus. As with each of the visual cortical areas, each of the visual dorsal thalamic (lateral geniculate, lateral posterior, posterior thalamic) nuclei relate to a separate territory of the reticular nucleus, with little or no overlap. Each of these dorsal thalamic territories within the reticular nucleus receives inputs from one or more of the visual cortical areas. For instance, the region of the reticular nucleus that is labelled after an injection into the lateral geniculate nucleus encompasses the reticular regions which receive afferents from cortical areas 17, rostral 18b and caudal 18b. These results suggest that individual cortical areas may influence the activity of different dorsal thalamic nuclei through their reticular connections.