Cortical visual areas of the cat project differentially onto the nuclei of the accessory optic system

Cortical projections to the nuclei of the cat's accessory optic system were demonstrated by anterograde transport and degeneration methods. Areas 21a, 21b, AMLS and PMLS were found to project to the medial, lateral and dorsal terminal accessory optic nuclei. Selective projections from area PLLS to the lateral terminal nucleus and from areas 17 and 18 to the medial terminal nucleus were noted. No terminal labeling was detected following injections of areas ALLS, DLS, 20a, 20b, 19, 7 or the splenial visual area. The accessory optic system has been implicated in the control of optokinetic nystagmus. Additional evidence supports a role for ipsilateral visual cortical projections in mediating optokinetic pursuit in the naso-temporal direction under monocular conditions. Thus the visual cortical projections we describe may partially underlie the observed functional laterality of monocularly elicited optokinetic pursuit in the cat. The present results further indicate that suprasylvian areas AMLS, PMLS and 21 are the cortical regions primarily responsible for descending visual influences on the cat's accessory optic nuclei.     

Corticopontine projections of the lateral suprasylvian cortex: De-emphasis of the central visual field

Layer V pyramidal cells of the cat lateral suprasylvian visual areas project to the pontine nuclei. Although all 6 of the suprasylvian visual areas project to the pons, the denset projections are from 3 areas: anterior medial lateral suprasylvian (AMLS), posterior medial lateral suprasylvian (PMLS) and ventral lateral suprasylvian (VLS). The organization of the corticopontine pathway from one of these areas (PMLS) suggests a disproportionate representation of the peripheral visual fields. This pattern of projection would serve to de-emphasize the central visual field.     

Thalamic projections to visual areas of the middle suprasylvian sulcus in the cat

The thalamic afferents to two areas of the lateral suprasylvian visual cortex in the cat were studied by using retrograde transport of horseradish peroxidase (HRP). Injections were localized retinotopically with electrophysiological recording. The posteromedial lateral suprasylvian area (PMLS) of Palmer et al. ('78) receives afferents from the pulvinar (P), the posterior nucleus of Rioch (PN), the C-laminae of the lateral geniculate nucleus (LGNd) and the centrolateral (CL), lateral posterior (LP), medial interlaminar (MIN) nuclei. The anteromedial lateral suprasylvian area (AMLS) receives afferents from CL, P, LP, PN, MIN, and probably from the posterior nuclear group (PO), and the lateral dorsal (LD) and ventral anterior (VA) nuclei. The LP-pulvinar complex has been divided into four zones on the basis of connectivity: geniculate wing, pulvinar, the lateral division of LP, and the interjacent division of LP (Updyke, '77; Graybiel and Berson, '80; Guillery et al., '80). The locations of labeled cells in the present experiments suggest that both AMLS and PMLS receive afferents from each of the four zones, although differences exist in the strength of the projections. While AMLS and PMLS receive afferents from many of the same nuclei (CL, P, LP, PN, and MIN), differences in their afferents also were noted. These differences are of three types. The first is that some nuclei project to only one of the cortical areas. PMLS alone receives input from the C-laminae of the LGNd while AMLS alone receives probable input from PO, LD, and VA. The second difference is in the strength of the projection from some nuclei. AMLS receives a stronger projection from CL and P than does PMLS. The third difference concerns the pattern of distribution of neurons that project to each cortical area. Labeled cells in LP are dispersed after an AMLS injection, but are found in clusters or bands after a PMLS injection. Thus our results indicate that the thalamic afferents to AMLS and PMLS are in general similar: however, differences in input to AMLS and PMLS suggest that inputs to PMLS are predominantly visual while AMLS receives a broader spectrum of afferent information.     

Cortical areas involved in horizontal OKN in cats: metabolic activity

Cerebral cortex improves optokinetic responses to high target velocities, but the specific cortical areas involved are unknown. Using the 14C-deoxyglucose technique, we compared local rates of cerebral glucose utilization in cats viewing a moving optokinetic nystagmus (OKN) drum (experimental group) with those in cats viewing a stationary OKN drum (control group). In the experimental group, glucose utilization was increased in areas 17 and 18 and in 4 areas in suprasylvian cortex (21a, 21b, PMLS, and VLS). There were no changes in glucose utilization in areas 7, 19, 20a, 20b, ALLS, AMLS, DLS, PLLS, the posterior suprasylvian area, and the splenial visual area. The increases in glucose utilization in areas 17 and 18 were most significant in the granular layers (inner III and IV). In areas 21a, 21b, PMLS, and VLS, the increases in glucose utilization extended from layers II through V. There was also a regional distribution of the increase in glucose utilization within each of these areas in the experimental animals. The increase in glucose utilization did not include the rostral portion of PMLS or the borders between areas PMLS and 21a, and VLS and 21b. In addition, there was a smaller increase in glucose utilization at the borders between areas 17 and 18 than in other portions of these 2 areas. The results indicate that areas 17, 18, 21a, 21b, PMLS, and VLS may be involved in the cortical modulation of horizontal OKN. The laminar distribution of label within the cortical areas corresponds with the distribution of projections from the dorsal lateral geniculate nucleus to areas 17 and 18, and from areas 17 and 18 to PMLS. The regional distribution of the metabolic activity within areas 17, 18, and PMLS coincides with that portion of cortex expected to be excited by either the spatial frequency of the stimulus or the retinalslip velocity (drum velocity minus slow phase eye velocity) occurring during the eye movements.     

Organization of visual corticostriatal projections in the cat,with observations on visual projections to claustrum and amygdala

Electrophysiological mapping criteria were employed to identify visual areas 20a, 20b, 21a, 21b, PMLS, AMLS, ALLS, PLLS, DLS, VLS, and PS in the cat, and to guide placement of tracer deposits. Anterograde tracer methods were used to study the corticostriatal projections of these extrastriate visual areas. The experiments demonstrate that all 11 extrastriate areas send projections to two distinct regions within the striatum, an extensive longitudinal zone within the caudate nucleus, and a more compact region within the posterolateral putamen. Cortical visual projections to the putamen terminate in relatively compact sheets or slabs, and appear to overlap extensively, while those to the caudate nucleus are irregularly patchy and more widely dispersed. Retrograde tracer deposits into the visual recipient zone of the caudate nucleus reveal substantial convergence of other cortical inputs to this same domain. Aspects of visuotopic organization are preserved in the visual projections to both the putamen and the caudate nucleus, but unequivocal retinotopic organization could not be inferred from the available material. Ten of the eleven extrastriate visual area also project topographically onto the visual zone of the claustrum. Area PS does not appear to contribute to the corticoclaustral projections. Five of the extrastriate visual areas (ALLS, PLLS, DLS, VLS, PS) also send sparse projections to the amygdaloid complex. c 1993 Wiley-Liss, Inc.     

Cortical and subcortical afferent connections of a posterior division of feline area 7 (area 7p)

Area 7 of the cat, as identified cytoarchitecturally, includes cortex both on the middle suprasylvian gyrus and on the anterior lateral gyrus. The aim of the experiments reported here was to determine whether within this zone there are subdivisions with qualitatively different patterns of afferent connectivity. Deposits of distinguishable retrograde tracers were placed at 29 sites in and around area 7 of 15 cats; cortical and subcortical telencephalic structures were then scanned for retrograde labeling. Our results indicate that cortex on the anterior lateral gyrus, although often included in area 7, is indistinguishable on connectional grounds from adjacent somesthetic cortex (area 5b). Cortex with strong links to visual, oculomotor, and association areas is confined to the middle suprasylvian gyrus and the adjacent lateral bank of the lateral sulcus. We refer to this discrete, connectionally defined zone as posterior area 7 (area 7p). Area 7p receives input from visual areas 19, 20a, 20b, 21a, 21b, AMLS, ALLS, and PLLS; from frontal oculomotor cortex (areas 6m and 6l); and from cortical association areas (posterior cingulate cortex, the granular insula, the posterior ectosylvian gyrus, and posterior area 35). Thalamic projections to area 7p arise from three specific nuclei (pulvinar; nucleus lateralis intermedius, pars caudalis; nucleus ventralis anterior) and from the intralaminar complex (nuclei centralis lateralis, paracentralis and centralis medialis). Neurons in a division of the claustrum immediately beneath the somatosensory and visual zones project to area 7p. Within area 7p, anterior-posterior regional differentiation is present, as indicated by the spatial ordering of projections from cingulate and frontal cortex, the thalamus, and the claustrum. Area 7p, as delineated by connectional analysis in this study, resembles cortex of the primate inferior parietal lobule both in its location relative to other cortical districts and in its pattern of neural connectivity.     

Corticotectal projections in the cat: anterograde transport studies of twenty-five cortical areas.

Retrograde transport studies have shown that widespread areas of the cerebral cortex project upon the superior colliculus. In order to explore the organization of these extensive projections, the anterograde autoradiographic method has been used to reveal the distribution and pattern of corticotectal projections arising from 25 cortical areas. In the majority of experiments, electrophysiological recording methods were used to characterize the visual representation and cortical area prior to injection of the tracer. Our findings reveal that seventeen of the 25 cortical areas project upon some portion of the superficial layers (stratum zonale, stratum griseum superficiale, and stratum opticum, SO). These cortical regions include areas 17, 18, 19, 20a, 20b, 21a, 21b, posterior suprasylvian area (PS), ventral lateral suprasylvian area (VLS), posteromedial lateral suprasylvian area (PMLS), anteromedial lateral suprasylvian area (AMLS), anterolateral lateral suprasylvian area (ALLS), posterolateral lateral suprasylvian area (PLLS), dorsolateral lateral suprasyvian area (DLS), periauditory cortex, cingulate cortex, and the visual portion of the anterior ectosylvian sulcus. While some of these corticotectal projections target all superficial laminae and sublaminae, others are more discretely organized in their laminar-sublaminar distribution. Only the corticotectal projections arising from areas 17 and 18 are exclusively related to the superficial layers. The remaining 15 pathways innervate both the superficial and intermediate and/or deep layers. The large intermediate gray layer (stratum griseum intermedium; SGI) receives projections from almost every cortical area; only areas 17 and 18 do not project ventral to SO. All corticotectal projections to SGI vary in their sublaminar distribution and in their specific pattern of termination. The majority of these projections are periodic, or patchy, and there are elaborate (double tier, bridges, or streamers) modes of distribution. We have attempted to place these findings into a conceptual framework that emphasizes that the SGI consists of sensory and motor domains, both of which contain a mosaic of connectionally distinct afferent compartments (Illing and Graybiel, '85, Neuroscience 14:455-482; Harting and Van Lieshout, '91, J. Comp. Neurol. 305:543-558). Corticotectal projections to the layers ventral to SGI, (stratum album intermediale, stratum griseum profundum, and stratum album profundum) arise from thirteen cortical areas. While an organizational plan of these deeper projections is not readily apparent, the distribution of several corticotectal inputs reveals some connectional parcellation.     

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 cortical and subcortical projections to area 6m of the cat

By analyzing regional variations of afferent connectivity, we have identified a medial subdivision of feline area 6 (area 6m) which differs from all surrounding sectors of the frontal lobe in its pattern of inputs. Area 6m is located in the ventral bank of the cruciate sulcus and on the adjacent medial face of the frontal lobe and is partially coextensive with the medial frontal eye field as identified previously in electrophysiological experiments. Area 6m is innervated by axons from visual, association, and oculomotor areas and does not receive projections from somesthetic or somatomotor areas. Cortical sources of input to area 6m include several retinotopically organized extrastriate visual areas (AMLS, ALLS, and PLLS), association areas with strong links to the visual system (area 7, granular insula, posterior ectosylvian gyrus, and cingulate gyrus), and a lateral division of area 6 (area 61) with oculomotor functions. Thalamic afferents of area 6m derive from the paralamellar ventral anterior nucleus, from a dorsolateral division of the mediodorsal nucleus, and from the rostral intralaminar nuclei. The claustrum and the basolateral nucleus of the amygdala project to area 6m. Projections from area 7, the posterior cingulate area, the ventral anterior nucleus, and the mediodorsal nucleus are spatially ordered in a pattern such that parts of area 6 close to the fundus of the cruciate sulcus receive input from neurons positioned anteriorly in the cortical areas, dorsolaterally in the ventral anterior nucleus, and ventrolaterally in the mediodorsal nucleus. Our results indicate that area 6m probably is involved in the voluntary control of gaze and attention rather than in skeletomotor functions.     

Graded classes of cortical connections: quantitative analyses of laminar projections to motion areas of cat extrastriate cortex

Current hierarchical models of the cerebral cortex are mainly based on qualitative connection studies. From wheatgerm-agglutinin-horseradish peroxidase injections, we examined the laminar patterns of projections to and between the three major subdivisions of the motion-processing lateral suprasylvian (LS) complex [areas posteromedial lateral suprasylvian area (PMLS), anteromedial lateral suprasylvian (AMLS), posterolateral lateral suprasylvian area (PLLS)] of cat extrastriate cortex and of an adjoining form-processing area, 21a. We counted approximately 145,000 labelled projection cells in 20 cortical areas in 11 cats, and applied various analyses to the data, expressed as the percent supragranular layer (%SG) origin of each connection. We report two main results. (i) A wide range of %SG values was obtained, both from each individual cat and across the 163 projections examined. Nonetheless, both hierarchical and non-parametric cluster analyses of the pooled connection origins suggested the presence of three distinct laminar projection classes, constrained by graded %SG values of 0-33%, 39-69% and 76-97%. These conformed, respectively, to feedback, lateral and feedforward laminar patterns seen qualitatively in our material. (ii) Hierarchical connectivity analyses suggested that PMLS, AMLS and PLLS are ordered in a hierarchical sequence. Macaque motion areas V5/MT, MST and FST are arranged in a similar sequence, and areas at equivalent levels of the two motion hierarchies have some analogous functional specializations. Our findings provide the first objective support for the three laminar projection classes that underpin existing theoretical models of hierarchical cortical organization, and they suggest that the implementation of higher-order motion processing evolved along similar lines in the cat and monkey visual cortex.     

Auditory activation of cortical visual areas in cats after early visual deprivation

Auditory activation of the primary visual cortex (area 17) and two extrastriate visual cortical areas - the anterolateral lateral suprasylvian area (ALLS) and anteromedial lateral suprasylvian area (AMLS), was investigated in visually impaired cats. Impairment was accomplished shortly after birth by bilateral eyelid suturing (binocularly deprived cats, BD) or bilateral enucleation (binocularly enucleated cats, BE). In BE cats, the optic nerve and chiasm were entirely degenerated. No cortical atrophy or cytoarchitectural malformation was noticed in either BD or BE cats. In both normal and impaired cats we found auditory-responsive cells in the ALLS and AMLS, areas that are considered strictly visual. The most remarkable finding was an increase in the relative number of these auditory cells in the BD and BE cats, which was more prominent in the latter. Some auditory-responsive cells were also found in area 17 of BE cats. On the basis of formal calculation, it is tempting to suggest that the increase in relative number of auditory cells in these areas reflects the transformation of all the visual cells in the ALLS of BD and BE cats into auditory cells. In BE cats, all bimodal cells and an appreciable percentage of non-responsive cells also had transformed to auditory cells. In the AMLS of BD cats, it is primarily the bimodal cells that become auditory cells, whereas in BE cats all the visual and bimodal cells as well as non-responsive cells undergo this transformation. This assumption, however, is one possible interpretation of our results but not the only one. Other modes of neuronal plasticity that might yield similar results in the visually deprived cats can not be ruled out.     

Divergence and collateral axon branching in subsystems of visual cortical projections from the cat lateral posterior nucleus.

The projections from the lateral (LPl), intermediate (LPi) and medial (LPm) subdivisions of the cat lateral posterior nucleus (n. LP) to visual areas 17, the posteomedial (PMLS) and posterolateral (PLLS) lateral suprasylvian and anterior ectosylvian (AEV) were studied using the retrograde labeling technique following concomitant injections of fluorescent dyes (Fast blue, Nuclear yellow, Evans blue and Rhodamine beta-isothiocyanate) into the different cortical loci. The results showed a medial-lateral topographical reversal of the visual n. LP-cortical connections: The ventral portion of LPl projects to area 17 whereas more dorsolateral regions of LPl and lateral LPi provide input to PMLS. Cells in medial LPi project mainly to the PLLS cortex and AEV receives afferents from the LPm. Areas of overlap were identified within the ventral LPl which projects to both area 17 and PMLS and within the LPi/LPm border region at the origin of connections to both PLLS and AEV. Furthermore, some single neurons within the areas of overlap were found to be double-labeled indicating divergent projections to their respective cortical targets via collateral axon branching. The data show that divergence and axonal branching are common features of the different n.LP-visual cortical subsystems and support the notion of the existence of families of thalamo-cortical systems which are distinct in their connection patterns and underlying functional properties.     

Afferent connections of the thalamic intralaminar nuclei in the cat

Afferents to the central lateral (CL), paracentral (PC) and central medial (CE) intralaminar nuclei (ILN) from cortical and subcortical sites were studied in the cat. We utilized stereotaxically guided injections of HRP into the CL and PC nuclei and tritiated leucine injections into various visual, parietal and limbic areas of cortex to demonstrate these connections. In studying the relatively weak visual cortical projections to the ILN, we demonstrated projections from areas 19, 20a, 21a, 21b, AMLS, PMLS and PLLS. However, our HRP injections into the ILN often revealed only a few labeled cells in any of the above areas; therefore conclusions regarding the absence of projections to ILN from remaining visual cortical areas should be made cautiously. The ILN receive heavier projections from the frontal eye fields, cingulate cortex, splenial cortex, insular cortex, somatosensory areas SI and SII, auditory areas SF, AII, and Ep, and parietal areas 5 and 7. The most robust projections appear to be from from frontal eye fields, cingulate and parietal areas. No topography was apparent in the projections to the ILN. All cortical projections originate ipsilaterally from layers V and VI. Heavy subcortical projections to the ILN originate in the pretectum, superior colliculus, reticular formation, and periaqueductal grey. Fewer afferents arise from several other brainstem and thalamic nuclei.     

Efferent projections of the thalamic intralaminar nuclei in the cat

Efferent projections of the central lateral (CL), paracentral (PC) and central medial (CE) intralaminar nuclei (ILN) to cortical and subcortical sites were studied in the cat. The combined methods of electrophysiologically guided cortical injections of tritiated leucine and stereotaxic injections of horseradish peroxidase (HRP) into the CL and PC nuclei were utilized. Additionally, fluorescent double-labeling techniques demonstrated patterns of intralaminar axon collateralization. We found that the ILN project ipsilaterally to all visual cortical areas except area 17. Projections to visual cortex are not arranged topographically or retinotopically. The ILN also project to the frontal eye fields (areas 6 and 8), anterior cingulate gyrus, suprasylvian fringe of the auditory cortex, insular cortex, parietal areas 5 and 7, caudate nucleus and claustrum. We noted especially heavy projections to the frontal eye fields and parietal areas 5 and 7. Fibers from the ILN terminate in cortical layers I and VI, and at the layer III-IV border. The demonstration of collateralization of ILN axons to two separate cortical areas implies that the same neuronal message may pass from the ILN to multiple cortical areas. It is concluded that the ILN may mediate a general cortical activation and may play a role in attention to visual, auditory and somatosensory (especially nociceptive) stimuli.     

Thalamocortical connections of the rostral intralaminar nuclei: an autoradiographic analysis in the cat

In this study the pattern of projections from the rostral intralaminar thalamic nuclei to the cerebral cortex was examined in the cat by autoradiography. Injections of tritiated proline and leucine were placed into the central lateral, paracentral, central medial, and para-stria medullaris nuclei. After injections into the central lateral nucleus, label is present on the lateral side within the presylvian sulcus, in most of the suprasylvian gyrus, including the adjacent lateral and suprasylvian sulci, and in the posterior corner of the ectosylvian gyrus. On the medial side, label is present in the orbitofrontal (Of), precentral agranular (Prag), anterior limbic (La), retrosplenial (Rs), and postsubicular (Ps) areas, as defined by Rose and Woolsey ('48a). The cingulate gyrus also contains label throughout (part of which was defined as the "cingular area," Cg, by Rose and Woolsey, '48a). Label is also found on both banks of the splenial and cruciate sulci. In addition, label is present within the lateral gyrus, on both its lateral and medial sides. The paracentral projections are similar to the central lateral input. On the lateral side, label is found within the presylvian sulcus, suprasylvian gyrus and adjacent lateral and suprasylvian sulci, and posterior ectosylvian gyrus. Medially, label is present in the Of, Prag, La, Cg, Rs, and Ps areas, and within the cruciate and splenial sulci, and in portions of the lateral gyrus. Following injections of the central medial nucleus, label is present in the presylvian sulcus; but in contrast to the central lateral and paracentral projections, the suprasylvian gyrus is labeled only in its posterior part. The central medial nucleus also projects to the posterior lateral gyrus, both laterally and medially. Also, the central medial nucleus projects heavily to rostral cortical zones, which include the Of, Prag and La areas, cruciate sulcus, and the rostral cingulate gyrus. The para-stria medullaris nucleus projects only to the presylvian sulcus and orbitofrontal cortex laterally, but, medially, has an extensive input similar to the central lateral and paracentral projections in that label is present in the Of, Prag, La, Cg, Rs, and Ps areas, in the cruciate and splenial sulci, and in the posterior lateral gyrus. The laminar distribution of label is as follows: the central lateral, paracentral and para-stria medullaris nuclei project primarily to layers I and III, whereas the central medial nucleus projects to layers I and VI. In addition, the central lateral projection has a patchy appearance in the retrosplenial and postsubicular cortices.     

Neurofilament protein: a selective marker for the architectonic parcellation of the visual cortex in adult cat brain.

In this immunocytochemical study, we examined the expression profile of neurofilament protein in the cat visual system. We have used SMI-32, a monoclonal antibody that recognizes a nonphosphorylated epitope on the medium- and high-molecular-weight subunits of neurofilament proteins. This antibody labels primarily the cell body and dendrites of pyramidal neurons in cortical layers III, V, and VI. Neurofilament protein-immunoreactive neurons were prominent in 20 visual cortical areas (areas 17, 18, 19, 20a, 20b, 21a, 21b, and 7; posteromedial lateral, posterolateral lateral, anteromedial lateral, anterolateral lateral, dorsal lateral, ventral lateral, and posterior suprasylvian areas; anterior ectosylvian, the splenial, the cingulate, and insular visual areas; and the anterolateral gyrus area). In addition, we have also found strong immunopositive cells in the A laminae of the dorsal part of the lateral geniculate nucleus (dLGN) and in the medial interlaminar nucleus, but no immunoreactive cells were present in the parvocellular C (1-3) laminae of the dLGN, in the ventral part of the LGN and in the perigeniculate nucleus. This SMI-32 antibody against neurofilament protein revealed a characteristic pattern of immunostaining in each visual area. The size, shape, intensity, and density of neurofilament protein-immunoreactive neurons and their dendritic arborization differed substantially across all visual areas. Moreover, it was also obvious that several visual areas showed differences in laminar distribution and that such profiles may be used to delineate various cortical areas. Therefore, the expression of neurofilament protein can be used as a specific marker to define areal patterns and topographic boundaries in the cat visual system.     

Organization of corticothalamic projections from parietal cortex in cat

Corticothalamic projections from areas 5a, 5b, and 7 of cat parietal cortex were studied with autoradiographic techniques. Each cortical area was identified by its cytoarchitectural characteristics and the patterns of termination were related to the thalamic nuclear groups. Injections of 3H-leucine in cortical area 5a were associated with terminal labeling primarily in the spinal recipient zone of the ventral lateral nucleus (VLsp) and the medial division of the posterior group (POm). The corticothalamic projections of area 5a are loosely topographically organized; medial parts of 5a project heavily to rostral and lateral parts of VLsp and sparsely to POm, while lateral parts of 5a project to more medial and caudal parts of VLsp and heavily to POm. Cortical area 5b projects primarily to the rostral portions of the lateral posterior nucleus (LP). These projections also appear to be topographically organized. The part of area 5b on the marginal gyrus projects to more ventral parts of rostral LP, while area 5b on the middle suprasylvian gyrus projects to more dorsal and lateral parts of rostral LP. Cortical area 7 projects to LP and the pulvinar (Pul). Rostral parts of area 7 project heavily to dorsal and lateral parts of LP and lightly to Pul; more caudal portions of area 7 projects relatively more heavily to Pul. The reticular, central lateral, and paracentral nuclei also receive projections, especially from the suprasylvian gyrus. The results are discussed with regard to putative sensory response characteristics of these cortical areas and to general thalamocortical organization.     

Cortical projections of the parvocellular laminae C of the dorsal lateral geniculate nucleus in the cat: An anterograde wheat germ agglutinin conjugated to horseradish peroxidase study

The areal and laminar distributions of the projection from the parvocellular part of laminae C of the dorsal lateral geniculate nucleus (Cparv) were studied in visual cortical areas of the cat with the anterograde tracing method by using wheat germ agglutinin conjugated to horseradish peroxidase. A particular objective of this study was to examine the central visual pathways of the W-cell system, the precise organization of which is still unknown. Because the Cparv in the cat is said to receive W-cell information exclusively from the retina and the superior colliculus, the results obtained would provide an anatomical substrate for the W-cell system organization in mammals. The results show that the cortical targets of the Cparv are areas 17, 18, 19, 20a, and 21a and the posteromedial lateral suprasylvian (PMLS) and ventral lateral suprasylvian(VLS) areas. In area 17, the projection fibers terminate in the superficial half of layer I; the lower two-thirds of layer III, extending to the superficial part of layer IV; and the deep part of layer IV, involving layer Va. These terminations form triple bands in area 17. The projection terminals in layer I are continuous, whereas those in layers III, IV, and Va distribute periodically, exhibiting a patchy appearance. In areas 18 and 19, the projection fibers terminate in the superficial half of layer I and in the full portions of layers III and IV, forming double bands. In these areas, the terminals in layer I are continuous, whereas those in layers III and IV distribute periodically, exhibiting a patchy appearance. In area 20a, area 21a, PMLS, and VLS, projection fibers terminate in the superficial part of layer I, in part of layer III, and in the full portion of layer IV, although they are far fewer in number than those seen in areas 17, 18, and 19. The present results demonstrate that the Cparv fibers terminate in a localized fashion in both the striate and the extrastriate cortical areas and that these W-cell projections are quite unique in their areal and laminar organization compared with the X- and Y-cell systems. J. Comp. Neurol. 392:439–457, 1998. © 1998 Wiley-Liss, Inc.     

The distribution of the cells of origin of callosal projections in cat visual cortex

The distribution of neurons projecting through the corpus callosum (callosal neurons) was examined in retinotopically defined areas of cat visual cortex. As many callosal neurons as possible were labeled in a single animal by surgically dividing the posterior two-thirds of the corpus callosum and exposing the cut ends of callosal axons to horseradish peroxidase. The distribution of callosal neurons within a visual field representation was related to standard electrophysiological maps as well as to recording sites marked by electrolytic lesions. Callosal neurons were found in every retinotopically defined cortical area. The portion of the visual field representation that contained callosal neurons increased progressively from the area 17/18 border to area 19, to areas 20 and 21, and to the lateral suprasylvian visual areas. In area 17, the portion of the visual field representation containing callosal neurons extended from the vertical meridian out to a maximum of 10 degrees azimuth. In the posteromedial lateral suprasylvian visual area, callosal neurons were present in a region extending from the vertical meridian representation out to a representation of 60 degrees azimuth. Most callosal neurons were medium to large pyramids at the border of layers III and IV. A few layer IV stellates were among the callosal neurons of areas 17 and 18. In area 19 and even more so in the lateral suprasylvian visual areas, callosal neurons included pyramidal and fusiform-shaped cells in layers V and VI. The laminar distributions of callosal neurons in areas 20 and 21 were similar to those of area 19 and the lateral suprasylvian visual areas. The widespread distribution of callosal neurons in areas 20 and 21 and in the lateral suprasylvian visual areas suggests that the regions of peripheral visual field representation in cat cortex, as well as the representations of the vertical meridian, have access to the opposite cerebral hemisphere. This finding is significant in light of demonstrations of the importance of some of these cortical areas in the interhemispheric transfer of visual learning.     

Projections from striate and extrastriate visual cortices of the cat to the reticular thalamic nucleus.

We have studied the pattern of connectivity of the visual cortical areas 17, 18, 19, 20a, 21a, posteromedial lateral (PMLS), and the posterolateral lateral (PLLS) suprasylvian areas with the reticular thalamic nucleus (RTN) of the cat ventral thalamus. Three cortical areas per hemisphere were injected iontophoretically with either 4% wheat germ agglutinin-horseradish peroxidase, 4% dextran-fluororuby, or 4% dextran-biotin. The visual field representations of the injection sites were determined by reference to previously published visuotopic maps of the cortex. The locations of labelled fibres, presumed terminals and cell bodies were determined with the aid of a camera lucida attachment and computer aided stereometry. In the ventral thalamus, the primary visual cortices (areas 17 and 18) project in a topographic manner to both the perigeniculate nucleus (PGN) and the RTN. By contrast, the "higher" visual cortical areas (areas 19, 21a, 20a, PMLS, and PLLS) project only to the RTN. Our experiments demonstrate the existence of a single, albeit coarse, visuotopic map within the RTN but do not support the notion of separate subregions within the RTN that can be related specifically to a particular visual cortical area. The putative single visuotopic map in the RTN appears to be organised in such a way that the vertical meridians are represented along the rostrocaudal axis of the RTN, whereas the horizontal meridians are mapped within the dorsoventral axis of the nucleus. The upper visual field is represented within regions of the RTN adjacent to the caudal part of the dorsal lateral geniculate nucleus (LGNd), whereas the lower visual field is represented in the parts of the RTN rostral to the LGNd. The map also shows a ventrodorsal shift along the rostrocaudal axis of the RTN such that in the rostral RTN the representation of vertical meridian is placed more ventrally than that in the caudal part of the nucleus.