Organization of the Visual Sector of the Thalamic Reticular Nucleus in Galago

Projections to and from the visual sector of the thalamic reticular nucleus were studied in the prosimian primate genus Galago by anterograde and retrograde transport of WGA-HRP injected into the dorsal lateral geniculate nucleus (GLd), pulvinar nucleus, and their cortical targets. Contrary to the idea that thalamic connections with the reticular nucleus are not delimited sharply between nuclei associated with the same modality, our results show a distinct laminar segregation of the projections from the GLd and pulvinar nuclei. The GLd is connected reciprocally with the lateral {frsol|2/3} of the caudal part of the reticular nucleus, and the striate cortex sends projections to the same lateral tier. Both sets of projections are organized topographically, lines of projection taking the form of slender elongated strips that run from caudo-dorsal to rostro-ventral within the nucleus. The pulvinar nucleus, which projects to several areas of the temporal, parietal, and occipital lobes, including the striate cortex, is connected reciprocally with the medial {frsol|1/3} of the caudal part of the reticular nucleus. Every injection into the pulvinar nucleus labelled a wide area of the medial tier, with no indication of visuotopic organization. The projections from the middle temporal area, one of the principal targets of the pulvinar nucleus, also terminate only in the medial tier of the visual sector. And we would expect that, in general, a thalamic nucleus and its cortical target would project to the same part of the reticular nucleus. The case of the striate area is an exception but only in the sense that it projects to the pulvinar nucleus as well as GLd. Thus an injection into a single locus in area 17 produces two parallel strips in the visual sector of the reticular nucleus, but both are in the lateral tier. We propose that each strip arises from a separate population of cells with cortical layer VI, one with an allegiance to the GLd and the other to the pulvinar nucleus.     

Pathways between striate cortex and subcortical regions in Macaca mulatta and Saimiri sciureus: Evidence for a reciprocal pulvinar connection

The efferent and afferent connections of striate cortex have been compared in Macaca mulatta and Saimiri sciureus monkeys after injecting striate cortex with a mixture of horseradish peroxidase and tritiated amino acids. Three reciprocal pathways were found in Macaca; they were between striate cortex and (i) all layers of the dorsal lateral geniculate nucleus; (ii) the inferior subdivision of pulvinar; and (iii) the lateral subdivision of pulvinar. In Saimiri the only recprocal pathway involved the dorsal lateral geniculate nucleus; the pulvinar received a heavy striate input but too few peroxidase-labeled neurons were found in pulvinar to demonstrate convincingly a projection to striate cortex. In both species the reticular and posterior nucleus of the thalamus and the superior colliculus receive striate input; in Saimiri the ventral lateral geniculate also receives a striate projection. In both species, neurons in the basal forebrain project to striate cortex. Striate cortex synaptic terminals in both dorsal lateral geniculate nucleus and pulvinar are topographically organized. Inferior and lateral subdivisions of pulvinar each contain a representation of central retina, and the fiber bundles separating the two subdivisions apparently mark the vertical meridian. The results of this experiment suggest that both retino-geniculate and retino-superior colliculus-pulvinar types of visual information may converge within the striate cortex.     

Thalamo-cortical projections in the tree shrew (Tupaia glis)

Cortical lesions were placed in 18 hemispheres, and thalamic degeneration was studied after a survival period of at least six weeks. Very small lesions within the striate area produced complete degeneration of neurons in a column through the lateral geniculate, from medial to lateral borders and comprising all of the laminae. Lesions of various loci within the striate area reveal a precise topographic projection, with the rostral lateral geniculate sending fibers to the caudal extremity of the striate area and the caudal lateral geniculate projecting to the rostral extremity of the striate; further, the dorso-ventral dimension in the lateral geniculate projects to the medio-lateral dimension in the striate area. Finally, the evidence from striate area lesions suggests that the lateral geniculate projections are confined to the striate area as defined by cytoarchitecture, which in turn corresponds precisely with visual area I as defined by electrophysiological recording. This conclusion is supported by the failure to find retrograde degeneration after lesions of the belt of cortex adjacent to the striate area. The temporal area which occupies an extensive section from V II to the rhinal fissure and the auditory cortex and which has been shown to be a visual receiving area, is the target of essential projections from the pulvinar. The pulvinar also sends sustaining collaterals within the temporal area and probably outside as well, especially to V II. However, the very crude topographic organization apparent in the pulvinar projections does not seem to be sufficiently refined to account for the organization of V II. A suggestion was made in closing that V II may be the result of convergent evolution in different mammalian lines of descent.     

Thalamic projections of the fusiform gyrus in man

Previous retrograde degeneration studies have shown that human extrastriate visual cortex receives projections from the pulvinar, but their precise topographical organization remained unknown. We report on the distribution of thalamic projections originating in the fusiform gyrus, as studied with the Nauta method for anterogradely degenerating axons, in a case of right fusiform gyrus infarction. Ipsilaterally to the lesion, high density of afferents was found in the inferior pulvinar nucleus and a low density in the medial pulvinar nucleus as well as in the postero-inferior part of the reticular nucleus; no degenerating fibres were found in the lateral geniculate body. Degenerating axons were completely absent in the contralateral thalamus. Thus, there is a precise topographic relationship between parts of the extrastriate cortex and the pulvinar, suggesting segregated thalamocortical pathways for different parts of the extrastriate cortex. As in nonhuman primates, the human inferior temporal cortex has no direct output to the lateral geniculate body.     

Subcortical projections, cortical associations, and some intrinsic interlaminar connections of the striate cortex in the squirrel monkey (Saimiri)

Laminar lesions made by the thermocoagulation in the lateral striate cortex of Saimiri reveal details of three distinct groups of descending fibers: interlaminar, cortico-cortical, and cortico-subcortical. The two most massive connections originate mainly from layer III. These are the interlaminar fibers terminating in the underlying layer V, and the systematically arranged projection upon area 18. Another cortical projection upon a narrow region in the superior temporal sulcus originates mainly from the infragranular layers, from which originate also the fibers passing to three subcortical structures, viz. the colliculus superior, the medial pulvinar and the griseum pontis. These subcortical projections are sparse compared to the interlaminar and cortico-cortical connections. In particular, no evidence was obtained that the lateral geniculate nucleus receives fibers from the lateral striate cortex. Lesions injuring the white matter, however, produced, in addition, anterograde degeneration in the nucleus praetectalis, three pulvinar nuclei, the lateral geniculate nucleus and the pregeniculate nucleus. This degeneration must have resulted from the interruption of fibers originating in areas other than the lateral striate cortex, and passing beneath the site of the lesion. The origin of the interlaminar fibers and of the association fibers upon area 18 provided strong evidence that, in Saimiri's striate cortex, the two sublayers traditionally referred to as sublayers IVa and IVb, actually form part of the conspicuously enlarged layer III which thus is subdivided into three distinct sublayers.     

Contribution of thalamic input to the specification of cytoarchitectonic cortical fields in the primate: Effects of bilateral enucleation in the fetal monkey on the boundaries,dimensions, and gyrification of striate and extrastriate cortex

Bilateral enucleation was performed at different fetal ages during corticogenesis, and the brains were prepared for histological examination. Early-enucleated fetuses (operated prior to embryonic day 77) showed morphological changes at the level of the thalamus and the cortex. In the thalamus, there was a loss of lamination and a decrease in size of the lateral geniculate nucleus. There was a decrease in the size of the inferior pulvinar, but there was no change in the lateral pulvinar. The border of striate cortex was as sharp in the enucleates as it was in the normal monkeys. In three of the four early enucleates, we observed an interdigitation of striate and extrastriate cortex. In three of the early enucleates, we observed a small island of nonstriate cortex near the striate border that was surrounded entirely by striate cortex. Enucleation led to an age-related reduction of striate cortex. This reduction was greater in the operculum than in the calcarine fissure. The reduction of striate cortex was accompanied by an increase in the dimensions of extrastriate visual cortex, so that the overall dimensions of the neocortex remained invariant. The extrastriate cortex in the enucleated animals presented a uniform cytoarchitecture and was indistinguishable from area 18 in the normal animal. There were changes in the gyral pattern that were restricted mainly to the cortex on the operculum. A deepening of minor dimples as well as the induction of a variable number of supplementary sulci led to an increase in the convolution of the occipital lobe. These results are discussed with respect to the specification of cortical areas. They demonstrate that the reduction in striate cortex was not accompanied by an equivalent reduction in the neocortex; rather, there was a border shift, and a large volume of cortex that was destined to become striate cortex appears to be cytoarchitectonically normal extrastriate cortex. © 1996 Wiley-Liss, Inc.     

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.     

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 visual cortex of the opossum: the retrograde transport of horseradish peroxidase to the lateral geniculate and lateral posterior nuclei.

The visual cortex of opossum was studied by injecting horseradish peroxidase into the cortex and identifying labeled neurons in the thalamus. The results show that the lateral geniculate nucleus projects to area 17 in a topographical manner: the rostral lateral geniculate is represented in caudal striate cortex, and the dorsal extremity of the lateral geniculate, which probably corresponds to the zero vertical meridian, is represented along the border of area 18. Small injections in area 17 produced restricted bands of labeled neurons across the medial-lateral extent of the lateral geniculate, suggesting a greater precision in the topography than previously shown by retrograde degeneration studies. Following injections into area 17, labeled cells were also found in the lateral posterior nucleus. Injections of peristriate cortex produced labeled cells in the lateral posterior nucleus, as well as the lateral intermediate, posterior and intralaminar nuclei. Since the lateral posterior nucleus receives visual projections from the superior colliculus, the results show two visual pathways: the geniculo-striate path projecting just to core area or area 17, and a more diffuse parallel path that projects to both the core and belt. Whether or not this overlap is characteristics of the mammalian prototype it seems to be present in widely separated species.     

A projection from the parabigeminal nucleus to the pulvinar nucleus in Galago.

A projection from the parabigeminal nucleus (Pbg) to the striate-recipient zone of the pulvinar nucleus in the prosimian Galago was identified by anterograde and retrograde transport methods. In addition to the pulvinar nucleus, Pbg projections were found to terminate in layers 4 and 5 of the dorsal lateral geniculate nucleus and the central lateral nucleus. All three of these structures project to the superficial layers of the striate cortex. Similarities between the Pbg in mammals and the nucleus isthmi in nonmammals in connections and neurochemistry reinforce the idea that these two nuclei are homologous.     

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.     

Visual cortex of the grey squirrel (Sciurus carolinensis): Architectonic subdivisions and connections from the visual thalamus

Thalamic projections to the visual cortex of the grey squirrel were studied by retrograde degeneration in the lateral geniculate and the pulvinar nuclei. The lateral geniculate was found to project to architectonic area 17 which also corresponds to visual area I as defined by its retinotopic organization. The projection is spatially organized in a precise way, and for every cortical point there is a corresponding column in the lateral geniculate which extends from border to border. For the binocular sector of area 17 the lateral geniculate column lies in the trilaminar part of the lateral geniculate, while for the uniocular sector the column lies in the bilaminar sector of the lateral geniculate. The pulvinar projects to several architectonic areas, areas 18 and 19 and to two or more temporal areas below area 19. This projection is roughly topographic but follows the sustaining pattern. When the squirrel is compared to the tree shrew and hedgehog what emerges is a conception of those changes in the visual system which arose as a result of adaptation to an arboreal habitat.     

Subcortical projections of area MT in the macaque

Area MT is a visuotopically organized area in extrastriate cortex of primates that appears to be specialized for the analysis of visual motion. To examine the full extent and topographic organization of the subcortical projections of MT in the macaque, we injected tritiated amino acids in five cynomolgus monkeys and processed the brains for autoradiography. The injection sites, which we identified electrophysiologically, ranged from the representation of central through peripheral vision in both the upper and lower visual fields and included, collectively, most of MT. Projections from MT to the superior colliculus are topographically organized and in register with projections from striate cortex to the colliculus. Unlike projections from striate cortex, those from MT are not limited to the upper layer of the stratum griseum superficiale but rather extend ventrally from the upper through the lower layer of the stratum griseum superficiale and even include the stratum opticum. Projections from MT to the pulvinar are organized into three separate fields. One field (P₁) is located primarily in the inferior pulvinar but extends into a portion of the adjacent lateral pulvinar. The second field (P₂) partially surrounds the first and is located entirely in the lateral pulvinar. The third and heaviest, projection field (P₃) is located posteromedially in the inferior pulvinar but also includes small portions of the lateral and medial pulvinar that lie dorsal to the brachium of the superior colliculus. While projections from MT to P₁ and P₂ are topographically organized, there appears to be a convergence of MT inputs to P₃. Projections from MT to the reticular nucleus of the thalamus are located in the ventral portion of the nucleus, approximately at the level of the caudal pulvinar. There was some evidence that MT sites representing central vision project more caudally than do those representing peripheral vision. Projections from MT to the caudate, putamen, and claustrum are localized to small, limited zones in each structure. Those to the caudate terminate within the most caudal portion of the body and the tail. Similarly, projections to the putamen are always to its most caudal portion, where the structure appears as nuclear islands. Projections to the claustrum are located ventrally, approximately at the level of the anterior part of the dorsal lateral geniculate nucleus. Projections from MT to the pons terminate rostrally in the dorsolateral nucleus, the lateral nucleus, and the dorsolateral portion of the peduncular nucleus. A topographic organization of these projections was not apparent, but there may be a heavier input from the part of MT representing peripheral vision than from the part representing central vision. The results indicate that while subcortical projections of MT in the macaque are more extensive than those of either striate cortex or V2, they are not more extensive than those of V4 and overlap them considerably. The lack of a unique set of subcortical projections from MT suggests that MT's contribution to subcortical visual processing lies in the unique information it supplies.     

Subcortical connections of area 17 in the tree shrew: an autoradiographic analysis

The present anterograde autoradiographic study reveals several targets of the striate cortex (area 17) of the tree shrew which were not previously observed in studies which used anterograde degeneration methods; our data also confirm several previous findings. The results are discussed in the context of these projections modulating ascending visual information (claustrum, lateral intermediate nucleus, pulvinar, dorsal lateral geniculate, cells of the external medullary lamina, reticular nucleus of the thalamus, superficial collicular layers, and the anterior and posterior pretectal nuclei) or visuomotor information (putamen, caudate, ventral lateral geniculate, pontine gray, and the anterior and posterior pretectal nuclei).     

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.     

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

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

Visuotopic organization of projections from striate cortex to inferior and lateral pulvinar in rhesus monkey

Anatomical material from two series of monkeys (Macaca mulatta)was used to determine the full extent and visuotopic organization of striate projections to the pulvinar. One series was processed for degeneration by the Fink-Heimer procedure following unilateral lesions of lateral, posterior, or medial striate cortex (representing the central, peripheral, and far peripheral visual field, respectively); collectively, the lesions included all of area 17. The second series was processed for autoradiography following tritiated amino-acid injections into striate sites representing the center of gaze and eccentricities ranging from 0.5° to greater than 30° from fixation in both the upper and lower fields. The results indicate the existence of two separate striate projection zones within the pulvinar. One, the PI/PL zone, is located primarily within the inferiorpulvinar (PI) but extends into the adjacentlateral pulvinar (PL). The other, the PL zone, is located entirely within the lateral pulvinar and partially surrounds the first zone along its dorsal, lateral, and ventral aspects. Within the PI/PL zone, striate projections are topographically organized and represent the entire contralateral visual field. Central vision is represented laterally and posteriorly, with the fovea represented at the caudal pole of the nucleus; conversely, far peripheral vision is found medially and anteriorly, adjacent to the medial geniculate nucleus. The representation of the horizontal meridian runs obliquely across PI/PL, such that the upper visual field is located ventrolaterally and the lower visual field dorsomedially. The representation of the vertical meridian is located along the lateral margin of PI in anterior sections of the pulvinar, but within PL in posterior sections. Thus, the vertical meridian appears to form the border between the lateral margin of the PI/PL zone and the medial margin of the PL zone. At the lateral margin of the PL zone is the representation of its horizontal meridian. Striate projections to the PL zone, unlike those to the PI/PL zone, are limited to the representation of central vision. These results suggest that striate inputs contribute to the visual properties of neurons (Bender, 1981 a) throughout the PI/PL zone, but are insufficient to explain the visual properties of neurons outside of the central visual field representation in the PL zone.     

Calbindin immunoreactivity in the geniculo-extrastriate system of the macaque: implications for heterogeneity in the koniocellular pathway and recovery from cortical damage

Although most projection neurons in the primate dorsal lateral geniculate nucleus (dLGN) target striate cortex (V1), a small number project instead to extrastriate visual areas and have been suggested to play a role in the preserved vision ("blindsight") that survives damage to V1. Moreover, the distribution of dLGN cells projecting to extrastriate bears a striking similarity to that of neurons that stain for calbindin D-28K (Cal), a calcium-binding protein involved in regulating neuronal excitability and considered a marker for the koniocellular or "K" pathway of geniculocortical processing. In these studies, we used double-labeling techniques to examine whether Cal content characterizes all or a subset of neurons making up the geniculo-extrastriate pathway in normal macaque monkeys. After injections of cholera toxin B-subunit into the prelunate gyrus, the proportion of retrogradely labeled neurons in the dLGN that were also immunoreactive for Cal varied from less than 40% to over 80%, indicating that only a subset of the geniculo-extrastriate projection falls within the K pathway as defined by Cal content. Analysis of the injected territories indicated that identity of the extrastriate cortical target may be systematically related to Cal content in the geniculo-extrastriate projection. To see whether the Cal-immunoreactive dLGN population might potentially play a role in preserved vision after V1 damage, we also examined the dLGN of a macaque that had sustained a lesion of V1 in infancy and survived until 4 years. In this animal, large, intensely Cal-immunoreactive neurons were found scattered throughout the otherwise degenerated dLGN zones and made up over 95% of the identifiable remaining neurons. The results support an emerging view that the macaque koniocellular system is highly heterogeneous in nature and also suggest that Cal content may be a critical feature of the pathway by which visual information reaches extrastriate cortex in the absence of V1.     

Ventral lateral geniculate nucleus projects to the dorsal lateral geniculate nucleus in the cat

The lamina C3 of the dorsal lateral geniculate nucleus of the cat does not receive retinal projections but instead receives visual information from the small subpopulation of W-type ganglion cells via the upper substratum of the stratum griseum superficiale of the superior colliculus. We herein report a projection from the lateral division of the ventral lateral geniculate nucleus into the lamina C3 of the dorsal lateral geniculate nucleus. As the lateral division receives projections from the contralateral retina and the ipsilateral upper stratum griseum superficiale of the superior colliculus, we suggest that these regions make up a small cell type W-cell neuronal network that provides visual information to layer I of the striate cortex via the lamina C3.     

Morphological and neurochemical comparisons between pulvinar and V1 projections to V2

The flow of visual information is clear at the earliest stages: the retina provides the driving (main signature) activity for the lateral geniculate nucleus (LGN), which in turn drives the primary visual cortex (V1). These driving pathways can be distinguished anatomically from other modulatory pathways that innervate LGN and V1. The path of visual information after V1, however, is less clear. There are two primary feedforward projections to the secondary visual cortex (V2), one from the lateral/inferior pulvinar and the other from V1. Because both lateral/inferior pulvinar and V2 cannot be driven visually following V1 removal, either or both of these inputs to V2 could be drivers. Retinogeniculate and geniculocortical projections are privileged over modulatory projections by their layer of termination, their bouton size, and the presence of vesicular glutamate transporter 2 (Vglut2) or parvalbumin (PV). It has been suggested that such properties might also distinguish drivers from modulators in extrastriate cortex. We tested this hypothesis by comparing lateral pulvinar to V2 and V1 to V2 projections with LGN to V1 projections. We found that V1 and lateral pulvinar projections to V2 are similar in that they target the same layers and lack PV. Projections from pulvinar to V2, however, bear a greater similarity to projections from LGN to V1 because of their larger boutons (measured at the same location in V2) and positive staining for Vglut2. These data lend support to the hypothesis that the pulvinar could act as a driver for V2. J. Comp. Neurol. 521:813–832, 2013. © 2012 Wiley Periodicals, Inc.