Evidence for Excitatory Amino Acid Neurotransmitters in Forward and Feedback Corticocortical Pathways within Rat Visual Cortex

It is a commonly accepted notion that cells which make projections between the multiple cortical areas found in the mammalian visual system are excitatory, but there is little direct evidence that this is the case. Here we demonstrate using retrograde tracing with D-[³H]aspartate that connections in the rat which project from lower to higher visual areas (i.e. forward) and those which project from higher to lower areas (i.e. feedback) may use excitatory amino acid neurotransmitters. Following injection into the primary visual cortex, clusters of retrogradely labelled cells were found in several extrastriate areas within the cytoarchitectonic subdivisions 18a (‘areas’ LM, AL, PX, FLX, RL, AX) and 18b (‘area’ MX), and in the retrosplenial cortex. In all of these areas D-[³H]aspartate-labelled cells were surrounded by diffuse label which may represent anterograde labelling of axon terminals. This suggests that both legs of reciprocal intracortical circuits have similar chemospecificity. To directly demonstrate excitatory amino acid localization in forward projections, D-[³H]aspartate was injected into extrastriate area LM. As expected, the results revealed retrogradely labelled neurons within area 17. Outside area 17, LM injections labelled neurons in AL, PX, FLX, RL, AX and MX. Taken in the context of the hierarchy of areas in rat cerebral cortex (Coogan and Burkhalter, J. Neurosci., 13, 3749–3772, 1993), these results show that D-[³H]aspartate labels: (1) forward connections from area 17 to LM, AL, PX, RL, AX and MX, (2) feedback connections from LM, AL, FLX, PX, RL, AX and MX to area 17, (3) feedback connections from AL, PX, RL, AX and MX to LM, and (4) lateral connections between FLX and LM. These findings strongly indicate that both forward and feedback connections as well as lateral connections at several different levels of the cortical hierarchy use excitatory amino acid neurotransmitters.     

Heterotopic and homotopic callosal connections in rat visual cortex

Heterotopic and homotopic callosal projections of rat visual cortex are evaluated. Callosal termination zones in visual cortex are identified with a degeneration technique following complete section of the corpus callosum. The zones which receive callosal afferents are the lateral one-third of area 17, an anteroposterior strip in dorsal and in ventral areas 18a, and 4 patches in area 18b. Following a large injection of lectin-bound horseradish peroxidase (WGA-HRP) into visual cortex, many retrogradely labeled neurons are found in the medial two-thirds of area 17 which does not receive callosal afferents, as well as in the lateral, callosal-recipient zone. These data suggest that heterotopic callosal pathways exist in visual cortex. Injections of tritiated amino acids into restricted parts of visual cortex show the following heterotopic connections: lateral area 17 projects to dorsal area 18a; medial area 17 projects to lateral area 17 and dorsal area 18a; area 18a projects to lateral area 17 and anteromedial area 18b; area 18b projects to lateral area 17, dorsal area 18a, heterotopic sites in area 18b, and to area 29d . Heterotopic connections are generally less dense than homotopic ones. In addition, heterotopic connections are generally less dense than homotopic ones. In addition, heterotopic projections terminate in the supragranular layers. This contrasts with the homotopic afferents of areas 17 and 18a which have additional strong projections to layer V. The distribution of label through the depth of the cortex in some of the callosal recipient zones has been quantified. Injections of WGA-HRP restricted to areas 17, 18a or 18b corroborate the presence of each of the heterotopic connections described above. Heterotopic afferents originate mostly from layer V neurons, whereas homotopic afferents arise from neurons primarily in layers II-V. Like the afferents, the numbers of callosal projection cells in heterotopic regions are substantially less than that in homotopic sites. heterotopic callosal connections may be one factor responsible for binocular vision and also may provide the basis for large, nonoriented receptive fields of units in layer V of rodent visual cortex.     

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.     

Distribution and morphological characterization of phosphate-activated glutaminase-immunoreactive neurons in cat visual cortex

Phosphate-activated glutaminase (PAG) is the major enzyme involved in the synthesis of the excitatory neurotransmitter glutamate in cortical neurons of the mammalian cerebral cortex. In this study, the distribution and morphology of glutamatergic neurons in cat visual cortex was monitored through immunocytochemistry for PAG. We first determined the specificity of the anti-rat brain PAG polyclonal antibody for cat brain PAG. We then examined the laminar expression profile and the phenotype of PAG-immunopositive neurons in area 17 and 18 of cat visual cortex. Neuronal cell bodies with moderate to intense PAG immunoreactivity were distributed throughout cortical layers II-VI and near the border with the white matter of both visual areas. The largest and most intensely labeled cells were mainly restricted to cortical layers III and V. Careful examination of the typology of PAG-immunoreactive cells based on the size and shape of the cell body together with the dendritic pattern indicated that the vast majority of these cells were pyramidal neurons. However, PAG immunoreactivity was also observed in a paucity of non-pyramidal neurons in cortical layers IV and VI of both visual areas. To further characterize the PAG-immunopositive neuronal population we performed double-stainings between PAG and three calcium-binding proteins, parvalbumin, calbindin and calretinin, to determine whether GABAergic non-pyramidal cells can express PAG, and neurofilament protein, a marker for a subset of pyramidal neurons in mammalian neocortex. We here present PAG as a neurochemical marker to map excitatory cortical neurons that use the amino acid glutamate as their neurotransmitter in cat visual cortex.     

Contribution of Area 17 to Cell Responses in the Striate-recipient Zone of the Cat's Lateral Posterior-Pulvinar Complex

The cat's lateral posterior-pulvinar complex (LP-pulvinar) contains three main representations of the visual field. The lateral part of the LP nucleus (LPI or striate-recipient zone) is the only region of these extrageniculate nuclei which receives afferents from the primary visual cortex. We investigated the contribution of area 17 to the response properties (orientation and spatial frequency tuning functions) of LPI neurons by cooling or lesioning the visual cortex. Responses of 40 LPI cells were studied before, during and after the reversible cooling of the striate cortex. When tested for orientation, a total of 10 units out of 28 was affected (36%). For most of these cells (eight of lo), cooling the visual cortex yielded a reduction of the cells’ visual responses without altering their orientation-selectivity (there was no significant change in the orientation tuning width). For only two cells, inactivation led to an increase in the response amplitude. Also, blocking the visual cortex never modified the direction-selectivity of LPI cells. When tested for spatial frequency, 12 neurons out of 33 were affected (36%) by the experimental protocol. In most cases, we observed a reduction in the responses at each spatial frequency tested, with no change in tuning bandwidth. For only three LPI cells, the effects of inactivation of the visual cortex were restricted to specific spatial frequencies, altering the profile of the spatial frequency tuning function. In five cats, removing area 17 reduced the proportion of visual neurons in LPI and the spared visually evoked responses were noticeably depressed. Despite the reduction in responsiveness, a few LPI receptive fields within the cortical scotoma were still sensitive to the orientation and/or direction of a moving stimulus. This last observation suggests that some properties in LPI could be generated either by circuits intrinsic to the LPI or by afferents from extrastriate cortical areas. Overall, these results indicate that projections from the visual cortex to the striate-recipient zone of the LP-pulvinar complex are mainly excitatory. Despite the strong impact of the area 17 projections, our data suggest that the extrastriate cortex could also play a role in the establishment of response properties in the cat's LPI.     

Afferents to a midbrain periaqueductal grey region involved in the ‘defence reaction’ in the cat as revealed by horseradish peroxidase. I. The telencephalon

Horseradish peroxidase injections were made at sites, within the midcollicular portion of the midbrain periaqueductal grey region (PAG), at which both electrical stimulation and subsequent microinjections of excitatory amino acids elicited defensive behaviour. Since excitatory amino acids depolarize cell bodies and dendrites located in the vicinity of the injection site but not axons of passage, the injections were centred within a PAG region known to contain neurones whose excitation elicited defensive behaviour. The telencephalic afferents to these sites were then determined. Sixty percent of the labelled telencephalic neurones were found in the frontal cortex, specifically in the medial frontal cortex along the banks of the rostral two-thirds of the cruciate sulcus, primarily area 6 and area 4, and the medial frontal cortex ventral to area 6 (area 32). Twenty-five percent of the labelled telencephalic neurones were found in the orbito-insular cortex while 8% were found in the parietal cortex surrounding the anterior ectosylvian sulcus. Although the functional significance of these projections remains to be established, available data suggest that these projections to the PAG arise from frontal 'oculomotor' and motor cortices, a polysensory insular cortical region and somatosensory, visual and auditory parietal cortical areas.     

Proportion of glutamate- and aspartate-immunoreactive neurons in the efferent pathways of the rat visual cortex varies according to the target.

Immunohistochemistry, with antisera directed against glutamate (Glu) or aspartate (Asp), was combined with wheat germ agglutinin-horseradish peroxidase (WGA-HRP) histochemistry to examine the distribution, morphology, and proportions of Glu- and Asp-containing neurons that give rise to corticofugal and callosal projections of the rat visual cortex. WGA-HRP injections in the dorsal lateral geniculate nucleus resulted in retrograde labelling of small and medium-sized cells throughout layer VI of the visual cortex. Of these cells, 60% were also Glu-immunoreactive and 61% Asp-positive. WGA-HRP injections in the superior colliculus labelled large and medium-sized neurons in the upper portion of layer V of the visual cortex. Of these cells, 46% were also stained for Glu and 66% for Asp. Injections in the pontine nuclei resulted in retrograde labelling of cells in the deeper part of cortical layer V. Retrogradely labelled cells, which were also immunoreactive for Glu or Asp, were large pyramidal cells. Corticopontine neurons, which were also Glu-positive, accounted for 42% of the total number of WGA-HRP labelled cells, whilst for Asp-positive neurons this percentage was 51%. Finally, after injections in the visual cortex, retrogradely labelled small and medium-sized cells were found throughout layers II-VI in the contralateral visual cortex. Of these neurons, 38% were also labelled for Glu while 49% were also Asp-immunoreactive. The present results demonstrate that substantial proportions of projection neurons in the rat visual cortex are immunoreactive for Glu or Asp, suggesting that these excitatory amino acids are the major transmitters used by the cortical efferent systems examined. Furthermore, the proportions of these immunoreactive neurons in the efferent pathways vary according to the target.     

In vivo microdialysis in the visual cortex of awake cat. II: sample analysis by microbore HPLC-electrochemical detection and capillary electrophoresis-laser-induced fluorescence detection

Sampling and monitoring release of excitatory and inhibitory amino acids in the striate cortex of mammals will provide important information for visual system research. Two microbore high performance liquid chromatography-electrochemical detection methods and a capillary electrophoresis-laser induced fluorescence detection were developed to determine the inhibitory amino acid, gamma-aminobutyric acid and the excitatory amino acids, glutamate and aspartate in microdialysates of cat striate cortex. In the liquid chromatography method, samples were derivatized using OPA-TBT. Ten microliters of derivatized product was injected onto the microbore column (100 x 1 mm i.d., C8) for quantitative analysis. Electrochemical detection was employed. In the capillary electrophoresis method, samples were derivatized using fluorescein isothiocyanate and separated in borate buffer within 15 min, then detected by a laser-induced fluorescence detector.     

The development of neuropeptide Y immunoreactive neurons in cat visual cortical areas.

The development of NPY-ir neurons and fibers in cat striate and extrastriate cortex was studied to determine whether temporal changes in the morphology, distribution and density of NPY-ir neurons during development would provide clues to the emergence of specific cortical areas. No differences in the number or distribution of NPY-ir neurons were observed at any age among the five visual cortical areas examined, area 17, 18, 19, posteromedial lateral suprasylvian and posterolateral lateral suprasylvian cortex. The number of NPY-ir neurons in cat visual cortical areas was higher in adult animals than in kittens. The proportion of NPY-ir neurons found in layer VIb was constant throughout life, suggesting that NPY immunoreactivity is not a marker for the transient neurons of the subplate. NPY-ir neuronal morphology was seen to 'flatten' in older animals with the development of sulci and increasing density of the brain. In contrast to the pattern observed with NPY-ir neurons, NPY-ir processes exhibited area-dependent differences during development. NPY-ir fibers grew into area 17 earlier and with a radial orientation which was not consistently observed laterally. This radial orientation was still apparent in adult brains in layer IV of area 17, though there was no orientation of fibers in the other laminae or other visual areas of the adult.     

Layering in lamina 6 of cat striate cortex

Retrograde transport after the injection of the tracer, wheatgerm agglutinin-horseradish peroxidase, into different neural sites revealed a layering of labelled cells in lamina 6 of the striate cortex of the cat. Depending on their destination, efferent cells were clustered at different levels in lamina 6 so that cells projecting to the claustrum congregated in the lower half of the lamina while those projecting to other parts of the visual cortex, in either ipsi- or contralateral hemispheres, were found principally in the upper half and the cells with axons passing to the lateral geniculate nucleus occupied the central expanse (middle three-fifths) of the lamina.     

The development of somatostatin immunoreactive neurons in cat visual cortical areas

The development of somatostatin immunoreactive (SOM-ir) neurons in cat striate and extrastriate cortex was studied to determine whether temporal changes in the morphology, distribution and density of SOM-ir neurons during development would provide clues to the emergence of specific cortical areas. The visual cortical areas examined included areas 17–19 and 7, posteromedial lateral suprasylvian, posterolateral lateral suprasylvian cortex and splenial visual area. We observed that the pattern of SOM-ir neurons in the cortical plate reflects the maturation of the cortical plate. At 1 week of age, SOM-ir neurons were only found in layers V and VI of the developing cortex; by 2 weeks of age, SOM-ir neurons were found in layer IV; and by 3 weeks of age, SOM-ir neurons were located in all layers of the cortex except layer I. SOM-ir neurons in the subplate were much more numerous under lateral cortical areas than under medial areas. This difference decreased over the first 2 postnatal weeks and by the 14th day after birth (P14), the distribution and numbers of SOM-ir neurons in the subplate/white matter had reached the adult pattern. The timing of exuberant SOM expression in the subplate suggests a function in the formation of visual corticocortical connections which begin to develop during the first postnatal week in the kitten.     

Electrophysiological and neurochemical study of the rat geniculo‐cortical pathway. Evidence for glutamatergic neurotransmission

The projection from the dorsal lateral geniculate nucleus to the primary visual cortex of the rat was studied electrophysiologically. Electrical stimulation of the dorsal lateral geniculate nucleus and the optic tract produced three types of responses on neurons of area 17: excitation followed by inhibition, excitation and inhibition. These results extend and confirm, in adult rats, previous studies done in rat geniculate-visual cortex cocultures preparations in vitro. The role of glutamate in the neurotransmission of the rat geniculo-cortical pathway was also investigated. In a first set of experiments, the effects of kynurenate, an antagonist of glutamate receptors, on visual cortex neurons with a monosynaptic excitatory response to dorsal lateral geniculate nucleus stimulation were studied. Microiontophoresis of kynurenate in area 17 neurons selectively suppressed the excitatory response to dorsal lateral geniculate nucleus and optic tract stimulation. In a second set of experiments, the effects of electrical stimulation of the dorsal lateral geniculate nucleus and the optic tract on the release of amino acids in the rat visual cortex in vivo were studied. Using the push–pull method, we perfused a discrete region of the visual cortex with artificial cerebrospinal fluid (CSF), and the amino acid content of the perfusates was analysed by high performance liquid chromatography (HPLC). Stimulation of either the dorsal lateral geniculate nucleus or the optic tract significantly increased glutamate release in area 17. The rest of the amino acids studied did not show significant changes. The results provide evidence for the participation of glutamate in the neurotransmission of the geniculo-cortical pathway in the rat.     

In vivo microdialysis in the visual cortex of awake cat. I: surgery, animal training and sampling

Sampling and monitoring release of excitatory and inhibitory amino acids in the striate cortex of mammals will provide important information for visual system research. A method allowing repeated microdialysis in the cortical layers of area 17 of the awake cat is described. Under visual control through a surgical microscope and using a stereotactic instrument, four probe guides are permanently implanted in area 17 of one hemisphere of the anesthetized animal and two fixation bars are mounted on the skull to allow fixation of the cat in a stereotactic frame. The implantation of four probe guides in the same hemisphere allows simultaneous sampling from different cortical regions subserving different parts of the visual field. A removable transparent cover protects the probe guides. After recovery from surgery the awake cats are trained to adapt to a fixation in a stereotaxic apparatus. Once adapted to that situation, the cats are used for 5 h in vivo microdialysis experiments without anesthesia.     

Laminar distribution of NMDA receptors in cat and monkey visual cortex visualized by [3H]-MK-801 binding

Glutamate is the major excitatory neurotransmitter of the mammalian central nervous system. Two major classes of glutamate receptors have been reported. The actions of glutamate on its N-methyl-D-aspartate (NMDA)-type receptor may underlie developmental and adult plasticity as well as neurotoxicity. The NMDA-type of glutamate receptor in cat and monkey visual cortex was visualized by means of in vitro receptor autoradiography with the noncompetitive NMDA-receptor antagonist [³H]-MK-801. The kinetics, performed on tissue sections, revealed an apparently single, saturable site with an approximate dissociation constant (KD) of 18.5 nM in cat and 15.9 nM in monkey visual cortex. Autoradiography, performed on frontal sections of cat and monkey visual cortex, revealed a heterogeneous laminar distribution of NMDA receptors. Cat areas 17,18,19, and the lateral suprasylvian areas exhibited a similar NMDA-receptor distribution. In these areas, NMDA receptors were most prominent in layer II and the upper part of layer III. In monkey striate cortex, NMDA receptors were primarily concentrated in layers II, upper III, IVc, V, and VI. In monkey secondary visual cortex, [³H]-MK-801 labeling was most prominent in layers II, V, and VI; whereas in the temporal visual areas included in this study layer II displayed the heaviest receptor labeling. In neither cat nor monkey could we observe significant differences in NMDA-receptor distribution between different retinotopic subdivisions within a single visual area. Neither did we detect any periodic changes in NMDA-receptor distribution that would correspond to the compartments defined by cytochrome-oxidase in monkey V1 and V2. © 1993 Wiley-Liss, Inc.     

Bihemispheric Collateralization of the Cortical and Subcortical Afferents to the Rat's Visual Cortex

A fluorescent dye (usually fast blue or rhodamine tagged latex microspheres) was injected into cortical area 17 (or area 17 and the lateral part of area 18b) of adult and juvenile (15 - 22 day old) Sprague-Dawley albino rats. Another fluorescent dye (usually diamidino yellow) was injected into cortical areas 17, 18a and 18b of the opposite hemisphere. The injections involved only the cortical grey matter. After postinjection survival of 2 - 14 days the distribution of retrogradely labelled mesencephalic and prosencephalic cells was analysed. Both small and large injections labelled retrogradely a substantial number of cells in specific and nonspecific dorsal thalamic nuclei (lateral geniculate, lateral posterior, ventromedial, several intralaminar nuclei and nucleus Reuniens) as well as a small number of cells in the preoptic area of the hypothalamus and the mesencephalic ventral tagmental area (VTA). While labelled thalamic cells contained only the dye injected into the ipsilateral cortex, a small proportion of hypothalamic and VTA cells was labelled with the dye injected into the contralateral cortex. Virtually none of the cells in these areas were double labelled with both dyes. Both small and large injections labelled cells in the ipsilateral telencephalic magnocellular nuclei of the basal forebrain and the caudal claustrum. A substantial minority of labelled cells in these structures was labelled by the dye injected into the contralateral cortex. Furthermore, a small proportion (about 1%) of claustral cells projecting to the ipsilateral cortex were double labelled with both dyes. In several cortical areas ipsilateral to the injected area 17, associational neurons were intermingled with commissural neurons projecting to the contralateral visual cortex. A substantial proportion of associational neurons projecting to ipsilateral area 17 also projected to the contralateral visual cortex (associational-commissural neurons). Thus, in visual area 18a, the associational-commissural neurons were located in all laminae, with the exception of lamina 1 and the bottom of lamina 6, and constituted about 30% of the neurons projecting to ipsilateral area 17. In paralimbic association area 35/13, associational-commissural neurons were located in lamina 5 and constituted about 20% of neurons projecting to ipsilateral area 17. In the limbic area 29d, the associational-commissural neurons were located in laminae 4, 5 and the upper part of lamina 6 and constituted about 10% of the associational-commissural neurons projecting to ipsilateral area 17. In oculomotor area 8, double-labelled neurons were located in lamina 5 and constituted about 10% of the neurons projecting to ipsilateral area 17. Thus, it appears that the axons of mesencephalic and diencephalic neurons projecting to the visual cortex do not send collaterals into both hemispheres. The bihemispheric projection to the rat's visual cortex originates almost exclusively in the retinotopically organized cortical area 18a and in integrative cortical areas 35/13, 29d and 8.     

Lack of Exuberance in Clustered Intrinsic Connections in the Striate Cortex of One-month-old Kitten

The topographies of the intrinsic tangential connections in layers II - IVab of one-month-old and adult cat's striate cortex were compared by plotting the distribution of retrogradely labelled neurons following injection of small amounts of wheat-germ agglutinin - horseradish peroxidase. In both cortices, the tracer reveals the characteristic cluster-like arrangement of intrinsic tangential connections. No differences were found between the two age groups in terms of either the numbers of cell clusters, their periodicities or their profiles, nor was there any difference in the lateral extent of the tangential connections. We conclude that one month postnatally, tangentially projecting intrinsic networks in the upper layers of the cat striate cortex lack exuberance and have an adult-like topography.     

Ectosylvian visual area of the cat: location, retinotopic organization, and connections

We have mapped out the ectosylvian visual area (EVA) of the cat in a series of single- and multiunit recording studies. EVA occupies 10-20 mm2 of cortex at the posterior end of the horizontal limb of the anterior ectosylvian sulcus. EVA borders on somatosensory cortex anteriorly, auditory cortex posteriorly, and nonresponsive cortex laterally. EVA exhibits limited retinotopic organization, as indicated by the fact that receptive fields shift gradually with tangential travel of the microelectrode through cortex. However, a point-to-point representation of the complete visual hemifield is not present. We have characterized the afferent and efferent connections of EVA by placing retrograde and anterograde tracer deposits in EVA and in other cortical visual areas. The strongest transcortical fiber projection to EVA arises in the lateral suprasylvian visual areas. Area 20, the granular insula, and perirhinal cortex provide additional sparse afferents. The projection from lateral suprasylvian cortex to EVA arises predominantly in layer 3 and terminates in layer 4. EVA projects reciprocally to all cortical areas from which it receives input. The projection from EVA to the lateral suprasylvian areas arises predominantly in layers 5 and 6 and terminates in layer 1. EVA is linked reciprocally to a thalamic zone encompassing the lateromedial-suprageniculate complex and the adjacent medial subdivision of the latero-posterior nucleus. We conclude that EVA is an exclusively visual area confined to the anterior ectosylvian sulcus and bounded by nonvisual cortex. EVA is distinguished from other visual areas by its physical isolation from those areas, by its lack of consistent global retinotopic organization, and by its placement at the end of a chain of areas through which information flows outward from the primary visual cortex.     

A comparison of the organization of the projections of the dorsal lateral geniculate nucleus, the inferior pulvinar and adjacent lateral pulvinar to primary visual cortex (area 17) in the macaque monkey

Both anterograde and retrograde transport tracing methods were used to study the organization of the projections of the dorsal lateral geniculate (DLG), the inferior pulvinar and subdivisions of the lateral pulvinar to primary visual cortex (striate cortex or area 17). The DLG projects only to striate cortex. These projections are retinotopically organized, and do not extend to any cortical layers above layer IVA. In contrast the inferior pulvinar (PI) and the immediately adjacent portion of the lateral pulvinar (PL alpha 48) project to both striate and prestriate cortex. The projections from these two thalamic areas to the striate cortex are also retinotopically organized and exist in parallel with those from the DLG. In contrast to the DLG, the projections from PI and PL alpha terminate above layer IVA in striate cortex, i.e. layers I, II and III. In prestriate cortex the layers of termination include layers IV, III and I. The pulvinar terminations in layers II and III of area 17 occur in segregated patches as do the geniculate terminations in layers IVC and IVA. On the other hand the pulvinar terminations in layer I which overlie those in layers II and III of area 17 appeared to be continuous. Control studies show that the remainder of the lateral pulvinar overlying PL alpha does not project to striate cortex. It is concluded that there are 3 visuotopically organized inputs from the lateral thalamus to primary visual cortex and that each of these inputs have different layers of termination. The inputs from PI and DLG can convey direct retinal inputs while those from PI and PL alpha can also be involved in intrinsic cortico-thalamocortical connection with prestriate cortex. It remains, then that it cannot be tacitly assumed that the ascending inputs which influence the response properties of the primary cortical neurons arise solely from the dorsal lateral geniculate nucleus. It is also argued that these inputs to the supragranular layers may be excitatory as those from the DLG to the IVth layer.     

Study of the afferent connections of the visual area of the lower bank of the cruciate sulcus of the cerebral cortex of the cat using the retrograde horseradish peroxidase axon transport technic

The possible visual inputs to the new visual area found in our experiments in the lower bank of cruciate sulcus of the cat cortex were studied with horseradish peroxidase technique. HRP was injected in the place preliminary identified as the visual area in physiological experiments with the same cats. Labelled neurons were found in the visual areas of the cortex (lateral suprasylvian and ectosylvian), in the parietal cortex (area 5 and 7) and in a small amount in the prefrontal and limbic cortex. In all experiments labelled neurons were found in the claustrum. In the thalamus labelled neurons were found in the nucleus medialis dorsalis, intralaminar nuclei (contralis lateralis, paracentralis and centralis medialis) and in the nuclei ventralis anterior, ventralis medialis, anteromedialis and reuniens. Some stained neurons were found in the midbrain in the stratum griseum centrale; the conclusion was made that the main sources of inputs to the studied area were different regions of the visual system or the structures of the brain closely connected with the latter.     

The contribution of the dorsal lateral geniculate nucleus to the total pattern of thalamic terminations in striate cortex of the Virginia opossum

These studies were carried out to show the manner of projection of the dorsal lateral geniculate nucleus and other thalamic nuclei to striate cortex in the Virginia opossum. In order to demonstrate these projections, lesions were made in the dorsal lateral geniculate nucleus, in the ventral lateral geniculate nucleus, in most of the thalamus on one side except for the dorsal lateral geniculate nucleus, and in the entire unilateral thalamus. Following various survival times, usually seven days, the brains were appropriately prepared and stained with procedure I of the Fink-Heimer technique. Dorsal lateral geniculate neurons project in a topographical manner only to certain layers of striate cortex. These projections from the lateral geniculate are compared with the same system in other mammals, and it is concluded that it is similar in all mammals studied, except for the cat. In the cat the lateral geniculate projects beyond the border of striate cortex, but even in the cat the layers of termination within striate cortex are apparently similar. The ventral lateral geniculate nucleus does not project to visual cortex. Dorsal thalamic nuclei other dian the lateral geniculate project to peristriate cortex and to layers VI and I of striate cortex. The finding that thalamic nuclei, other than the lateral geniculate nucleus, project to striate cortex has never been described as part of the visual pathways in other mammals. It is suggested that these additional projections arise mainly from the lateral nuclear group of the thalamus in the opossum, and must be considered in relation to any response characteristics and organization of striate cells determined from physiological studies. These multiple thalamic projections can provide the substrate for more than one representation or “map” of sensory information in striate cortex.