Neuronal projections from V1 to V2 in amblyopia

The mechanism of amblyopia in children with congenital cataract is not understood fully, but studies in macaques have shown that geniculate synapses are lost in striate cortex (V1). To search for other projection abnormalities in amblyopia, the pathway from V1 to V2 was examined using a triple-label technique in three animals raised with monocular suture. [(3)H]proline was injected into one eye to label the ocular dominance columns. Cholera toxin B subunit conjugated to gold (CTB-Au) was injected into V2 to label V1 projection neurons. Alternate sections were processed for cytochrome oxidase (CO) and CTB-Au, or dipped for autoradiography. Eight fields of CTB-Au-labeled cells in V1 opposite injection sites were plotted in layers 2/3 or 4B. After thin stripe injection, labeled cells were concentrated in CO patches. Despite column shrinkage, cells in deprived and normal columns were equal in size and density in both layers 2/3 and 4B. After pale or thick stripe injection, labeled cells were concentrated in interpatches. Only 23% of projection neurons originated from deprived columns. This reduction exceeded the degree of column shrinkage, a result explained by the fact that column shrinkage causes disproportionate loss of interpatch territory. These data indicate that early monocular form deprivation does not alter the segregation of patch and interpatch pathways to V2 stripes or cause selective loss or atrophy of V1 projection neurons. The effect of shrinkage of geniculocortical afferents in layer 4C following visual deprivation is not amplified further by attenuation of the amblyopic eye's projections from V1 to V2.     

Cortical and subcortical connections of V1 and V2 in early postnatal macaque monkeys

Connections of primary (V1) and secondary (V2) visual areas were revealed in macaque monkeys ranging in age from 2 to 16 weeks by injecting small amounts of cholera toxin subunit B (CTB). Cortex was flattened and cut parallel to the surface to reveal injection sites, patterns of labeled cells, and patterns of cytochrome oxidase (CO) staining. Projections from the lateral geniculate nucleus and pulvinar to V1 were present at 4 weeks of age, as were pulvinar projections to thin and thick CO stripes in V2. Injections into V1 in 4- and 8-week-old monkeys labeled neurons in V2, V3, middle temporal area (MT), and dorsolateral area (DL)/V4. Within V1 and V2, labeled neurons were densely distributed around the injection sites, but formed patches at distances away from injection sites. Injections into V2 labeled neurons in V1, V3, DL/V4, and MT of monkeys 2-, 4-, and 8-weeks of age. Injections in thin stripes of V2 preferentially labeled neurons in other V2 thin stripes and neurons in the CO blob regions of V1. A likely thick stripe injection in V2 at 4 weeks of age labeled neurons around blobs. Most labeled neurons in V1 were in superficial cortical layers after V2 injections, and in deep layers of other areas. Although these features of adult V1 and V2 connectivity were in place as early as 2 postnatal weeks, labeled cells in V1 and V2 became more restricted to preferred CO compartments after 2 weeks of age.     

Effects of experimental strabismus on the architecture of macaque monkey striate cortex

Strabismus, a misalignment of the eyes, results in a loss of binocular visual function in humans. The effects are similar in monkeys, where a loss of binocular convergence onto single cortical neurons is always found. Changes in the anatomical organization of primary visual cortex (V1) may be associated with these physiological deficits, yet few have been reported. We examined the distributions of several anatomical markers in V1 of two experimentally strabismic Macaca nemestrina monkeys. Staining patterns in tangential sections were related to the ocular dominance (OD) column structure as deduced from cytochrome oxidase (CO) staining. CO staining appears roughly normal in the superficial layers, but in layer 4C, one eye's columns were pale. Thin, dark stripes falling near OD column borders are evident in Nissl-stained sections in all layers and in immunoreactivity for calbindin, especially in layers 3 and 4B. The monoclonal antibody SMI32, which labels a neurofilament protein found in pyramidal cells, is reduced in one eye's columns and absent at OD column borders. The pale SMI32 columns are those that are dark with CO in layer 4. Gallyas staining for myelin reveals thin stripes through layers 2-5; the dark stripes fall at OD column centers. All these changes appear to be related to the loss of binocularity in cortical neurons, which has its most profound effects near OD column borders.     

Feedforward and feedback connections and their relation to the cytox modules of V2 in Cebus monkeys

To study the circuitry related to the ventral stream of visual information processing and its relation to the cytochrome oxidase (CytOx) modules in visual area V2, we injected anterograde and retrograde cholera toxin subunit B (CTb) tracer into nine sites in area V4 in five Cebus apella monkeys. The injection site locations ranged from 2° to 10° eccentricity in the lower visual field representation of V4. Alternate cortical sections, cut tangentially to the pial surface or in the coronal plane, were stained for CTb immunocytochemistry or for CytOx histochemistry or for Nissl. Our results indicate that the V4‐projecting cells and terminal‐like labeling were located in interstripes and thin CytOx‐rich stripes and avoided the CytOx‐rich thick stripes in V2. The feedforward projecting cell bodies in V2 were primarily located in the supragranular layers and sparsely located in the infragranular layers, whereas the feedback projections (i.e., the terminal‐like labels) were located in the supra‐ and infragranular layers. V4 injections of CTb resulted in labeling of the thin stripes and interstripes of V2 and provided an efficient method of distinguishing the V2 modules that were related to the ventral stream from the CytOx‐rich thick stripes, related to the dorsal stream. In V2, there was a significant heterogeneity in the distribution of projections: feedforward projections were located in CytOx‐rich thin stripes and in the CytOx‐poor interstripes, whereas the feedback projections were more abundant in the thin stripes than in the interstripes. J. Comp. Neurol. 522:3091–3105, 2014. © 2014 Wiley Periodicals, Inc.     

Projections between visual cortex and pulvinar in the rat

The extrageniculate visual pathway, which carries visual information from the retina through the superficial layers of the superior colliculus and the pulvinar, is poorly understood. The pulvinar is thought to modulate information flow between cortical areas, and has been implicated in cognitive tasks like directing visually guided actions. In order to better understand the underlying circuitry, we performed retrograde injections of modified rabies virus in the visual cortex and pulvinar of the Long‐Evans rat. We found a relatively small population of cells projecting to primary visual cortex (V1), compared to a much larger population projecting to higher visual cortex. Reciprocal corticothalamic projections showed a similar result, implying that pulvinar does not play as big a role in directly modulating rodent V1 activity as previously thought.     

Cortical projections to the two retinotopic maps of primate pulvinar are distinct

Comprised of at least five distinct nuclei, the pulvinar complex of primates includes two large visually driven nuclei; one in the dorsal (lateral) pulvinar and one in the ventral (inferior) pulvinar, that contain similar retinotopic representations of the contralateral visual hemifield. Both nuclei also appear to have similar connections with areas of visual cortex. Here we determined the cortical connections of these two nuclei in galagos, members of the stepsirrhine primate radiation, to see if the nuclei differed in ways that could support differences in function. Injections of different retrograde tracers in each nucleus produced similar patterns of labeled neurons, predominately in layer 6 of V1, V2, V3, MT, regions of temporal cortex, and other visual areas. More complete labeling of neurons with a modified rabies virus identified these neurons as pyramidal cells with apical dendrites extending into superficial cortical layers. Importantly, the distributions of cortical neurons projecting to each of the two nuclei were highly overlapping, but formed separate populations. Sparse populations of double-labeled neurons were found in both V1 and V2 but were very low in number (<0.1%). Finally, the labeled cortical neurons were predominately in layer 6, and layer 5 neurons were labeled only in extrastriate areas. Terminations of pulvinar projections to area 17 was largely in superficial cortical layers, especially layer 1.     

Tangential organization of callosal connectivity in the cat's visual cortex

Cells and/or terminals of corticocortical pathways in mammalian visual cortex often have a discontinuous distribution across the surface of the cortex. A modular organization of cortical function has been shown to underlie the tangential segregation of many inputs and outputs. Here, we present evidence that the callosal pathway in the visual cortex of the cat follows these general principles. Large injections of wheat germ agglutinin-horseradish peroxidase or biotinylated dextran amine were made in areas 17 and 18, and callosal labeling was analyzed in tangential sections. The band of callosal cells and terminals straddling the border of areas 17 and 18 was not uniform but varied in density in a complicated fashion. Fluctuations in density of callosal connections became more clear 2–3 mm lateral or medial to the 17/18 border, as the callosal labeling became less dense. Here, regular fluctuations with a periodicity of about 1 mm in area 17, and slightly greater than 1 mm in area 18 were apparent. Cytochrome oxidase staining in areas 17 and 18 showed a pattern of dense blobs with the same spacing as the callosal labeling in these areas, and the blobs were found to align with the patches of callosal labeling. Larger, more irregularly spaced stripes of callosal labeling extended from the lateral part of area 18 across area 19 and into more lateral visual areas. These results suggest that the callosal pathway in the cat's visual cortex has a patchy distribution similar to many ipsilateral corticocortical projections, and that the columnar system marked by cytochrome oxidase is important for the organization of (interhemispheric) corticocortical connectivity in cats. © 1994 Wiley-Liss, Inc.     

Multiple zonal projections of the nucleus reticularis tegmenti pontis to the cerebellar cortex of the rat

Compartmentalization (alternating labelled and unlabelled stripes) of mossy fibre terminals was found in the cerebellar cortex after iontophoretic injections of biotinylated dextran amine into discrete regions of the nucleus reticularis tegmenti pontis (NRTP). The zonal pattern was only observed when volumes of nuclear tissue ranging from 4.5 x 106 to 17.66 x 106 microm3 were impregnated. Up to nine compartments (i.e. up to five stripes separated by four interstripes) were found in crus I and in vermal lobule VI. Up to seven compartments (four stripes and three interstripes) were found in crus II; up to five compartments (three stripes and two interstripes) were identified in the lobulus simplex, the paraflocculus and vermal lobules IV, V and VII; up to three compartments (two stripes and one interstripe) were identified in the paramedian lobule and, finally, up to two compartments (one stripe and one interstripe) were identified in the copula pyramidis, in the flocculus and in vermal lobules II, III, VIII and IX. The projections of the NRTP are arranged according to a divergent/convergent projection pattern. From single injections in the NRTP, projections were traced to a set of cortical stripes widely distributed over the cerebellar cortex. The set of stripes labelled from different regions of the NRTP partially overlapped but complete overlap was never found. This finding revealed that the topographic combination of the projections of the NRTP to the cerebellar cortex is specific for each region of the NRTP. Finally, the projections to single cortical areas were arranged according to a pattern of compartmentalization that is specific for each cortical area, independent of the site of injection in the NRTP and of the number of stripes evident in the cortex.     

The visual pulvinar in tree shrews II. Projections of four nuclei to areas of visual cortex

Patterns of thalamocortical connections were related to architectonically defined subdivisions of the pulvinar complex and the dorsolateral geniculate nucleus (LGN) in tree shrews (Tupaia belangeri). Tree shrews are of special interest because they are considered close relatives of primates, and they have a highly developed visual system. Several distinguishable tracers were injected within and across cortical visual areas in individual tree shrews in order to reveal retinotopic patterns and cortical targets of subdivisions of the pulvinar. The results indicate that each of the three architectonic regions of the pulvinar has a distinctive pattern of cortical connections and that one of these divisions is further divided into two regions with different patterns of connections. Two of the pulvinar nuclei have similar retinotopic patterns of projections to caudal visual cortex. The large central nucleus of the pulvinar (Pc) projects to the first and second visual areas, V1 and V2, and an adjoining temporal dorsal area (TD) in retinotopic patterns indicating that the upper visual quadrant is represented dorsal to the lower quadrant in Pc. The smaller ventral nucleus (Pv) which stains darkly for the Cat-301 antigen, projects to these same cortical areas, with a retinotopic pattern. Pv also projects to a temporal anterior area, TA. The dorsal nucleus (Pd), which densely expresses AChE, projects to posterior and ventral areas of temporal extrastriate cortex, areas TP and TPI. A posterior nucleus, Pp, projects to anterior areas TAL and TI, of the temporal lobe, as well as TPI. Injections in different cortical areas as much as 6 mm apart labeled overlapping zones in Pp and double-labeled some cells. These results indicate that the visual pulvinar of tree shrews contains at least four functionally distinct subdivisions, or nuclei. In addition, the cortical injections revealed that the LGN projects topographically and densely to V1 and that a significant number of LGN neurons project to V2 and TD.     

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.     

Cortical connections of striate and extrastriate visual areas in tree shrews

The ipsilateral and contralateral cortical connections of visual cortex of tree shrews (Tupaia belangeri)were investigated by placing restricted injections of fluorochrome tracers, wheat germ agglutinin-horseradish peroxidase, or biotinylated dextran amine into area 17 (V1), area 18 (V2), or the adjoining temporal dorsal area (TD). As previously reported, V1 was characterized by a widespread, patchy pattern of intrinsic connections; ipsilateral connections with V2, TD, and to a lesser extent, other areas of the temporal cortex; and contralateral connections with V1, V2, and TD. A surface-view of the myelin pattern in V1 revealed a patchwork of light and dark module-like regions. The ipsilateral connections with V2 and TD were roughly topographic, whereas heterotopic locations in V1 were callosally connected. Injections in V2 labeled as much as one third of V2 in a patchy pattern, and portions of ipsilateral V1 and TD in roughly topographic patterns. In addition, connections with several other visual areas in the temporal lobe were revealed. Contralaterally, most of the label was in V2, with some in V1 and TD. Injections in TD demonstrated connections within the region, and with adjoining portions of the temporal cortex, V2, and V1. There were sparse connections with an oval of densely myelinated cortex, which we have termed the temporal inferior area (TI). Callosal connections were concentrated in TD, but also included V2. The results provide further evidence for modular organizations within V1 and V2, and reveal for the first time the complete patterns of cortical connections of V2 and TD. The results are consistent with the proposal that at least three visual areas, the temporal anterior area, TA, the temporal dorsal area, TD, and the temporal posterior area, TP, exist along the rostrolateral border of V2 in tree shrews; suggest visual involvement of at least three other areas, the temporal inferior area, TI, the temporal anterior lateral area, and the temporal posterior inferior area located more ventrally in the temporal cortex; and fortify the conclusion that TD is the likely homologue of the middle temporal visual area of primates. Because tree shrews are considered close relatives of primates, the evidence for several visual areas along the border of V2 is more compatible with theories that propose a series of visual areas along V2 in primates, rather than a single visual area, V3. J. Comp. Neurol. 401:109–128, 1998. © 1998 Wiley-Liss, Inc.     

Lack of robust LGN label following transneuronal rabies virus injections into macaque area V4

In primates, retinal inputs are relayed through the magno- and parvocells of the lateral geniculate nucleus (LGN) indirectly to extrastriate visual cortex. The most direct pathway identified to the extrastriate cortex is a disynaptic one that provides robust magno- and parvocellular inputs to the middle temporal area (MT). The inclusion of parvocells in this projection is somewhat surprising because of their importance for color and form vision, whereas MT is more strictly tuned to velocity. This raises the question of whether areas more involved in color and form processing, such as V4, receive similar projections. We report here on experiments that use rabies virus injections into V4 to retrogradely label mono- and disynaptic inputs. We find only a small number of labeled neurons in the LGN in a pattern consistent with monosynaptic labeling of koniocells, rather than disynaptic labeling of magno- and parvocells. The lack of robust magno- and parvocellular label was not due to ineffective viral transport because in the same cases we find hundreds of neurons labeled in the thalamic reticular nucleus, a structure that can only be labeled disynaptically from the cortex. We also find a complete absence of neurons labeled in V1, but thousands in adjacent areas V2 and V3. This result helps explain the absence of labeled magno- and parvocells in the LGN because disynaptic transport from an extrastriate visual area should require a relay through V1. Taken together, these results suggest that ascending magno/parvocellular inputs to V4 are more hierarchically organized than the relatively direct inputs to MT.     

Distribution of neurons projecting to the superior colliculus correlates with thick cytochrome oxidase stripes in macaque visual area V2

In visual area V2 of macaque monkeys, cytochrome oxidase (CO) histochemistry reveals a pattern of alternating densely labeled thick and thin stripe compartments and lightly labeled interstripe compartments. This modular organization has been associated with functionally separate pathways in the visual system. We examined this idea further by comparing the pattern of CO stripes with the distribution of neurons in V2 that project to the superior colliculus. Visually evoked activity in the superior colliculus is known to be greatly reduced by blocking magnocellular but not parvocellular layers of the lateral geniculate nucleus (LGN). From previous evidence that V2 thick stripes are closely associated with the magnocellular LGN pathway, we predicted that a significant proportion of V2 neurons projecting to the superior colliculus would reside in the thick stripes. To test this prediction, the tangential distribution of retrogradely labeled corticotectal cells in V2 was compared with the pattern of CO stripes. We found that neurons projecting to the superior colliculus accumulated preferentially into band-like clusters that were in alignment with alternate CO dense stripes. These stripes were identified as thick stripes on the basis of their physical appearance and/or by their affinity to the monoclonal antibody Cat-301. A significantly smaller proportion of labeled cells was observed in thin and interstripe compartments. These data provide further evidence that the spatial distribution of subcortically projecting neurons can correlate with the internal modular organization of visual areas. Moreover, they support the notion that CO compartments in V2 are associated with functionally different pathways. J. Comp. Neurol. 377:313–323, 1997. © 1997 Wiley-Liss, Inc.     

Thalamic inputs to visual areas 1 and 2 in the rabbit

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

Subcortical projections of six visual cortical areas in the owl monkey, Aotus trivirgatus.

Subcortical projections of six visual cortical areas (Areas 17 and 18, the Middle Temporal, Dorsomedial and Medial Areas, and the Posterior Parietal Region) in the owl monkey, Aotus trivigatus, were investigated with autoradiographic methods following injections of tritiated proline. No contralateral projections were demonstrated. While some brainstem structures received input from all six subdivisions of cortex, each cortical area appeared to exhibit its own unique pattern of subcortical projections. All six cortical areas were found to project to the superior and inferior divisions of the pulvinar, reticular nucleus of the thalamus, pretectum and superior colliculus. Other subcortical targets of one or more visual cortical areas were the basal ganglia, claustrum, zona incerta, one or more of the intralaminar nuclei, lateral posterior nucleus, pregeniculate nucleus, dorsal lateral geniculate nucleus, and pontine nuclei. Furthermore, details of corticofugal projections to the dorsal lateral geniculate nucleus, pretectum and superior colliculus varied with the cortical area studied. The projections to the reticular nucleus, pregeniculate nucleus, dorsal lateral geniculate nucleus, the inferior and a portion of the superior division of the pulvinar and the superior colliculus were found to be topographically organized. The targets of the subcortical projections were compared with those of the retina, as revealed by autoradiographic methods following tritiated proline injections of the eye and were found to overlap to varying extents in the superior colliculus, pretectum and dorsal lateral geniculate nucleus and to be segregated in the pregeniculate nucleus. The results substantiate the validity of previous studies in the owl monkey that suggest that the visual cortex is subdivided into several functionally distinct areas; and illustrate the complexity of corticofugal influence on visual processing.     

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.     

New view of the organization of the pulvinar nucleus in Tupaia as revealed by tectopulvinar and pulvinar-cortical projections

The projections of the superficial layers of the superior colliculus to the pulvinar nucleus in Tupaia were reexamined by injecting WGA-HRP into the tectum. The main result was finding two different patterns of terminations in the pulvinar nucleus: a zone remote from the lateral geniculate nucleus, which occupies the dorsomedial and caudal poles of the pulvinar nucleus, was almost entirely filled with terminals in every case irrespective of the location of the injection site; and a second division of the pulvinar nucleus, adjacent to the lateral geniculate nucleus, contained irregular patches--much more densely populated--and the distribution of patches varied from case to case. We call the first projection "diffuse" and the patchy projection "specific." Next we injected several divisions of the extrastriate visual cortex to find the cortical target of each pathway. The diffuse path terminates in the ventral temporal area (Tv). The specific path terminates in the dorsal temporal area (Td) and area 18. We speculated about the significance of the two pathways: the specific path may be responsible for the preservation of vision after removal of the striate cortex; the diffuse path may have an important place in the evolution of the visual areas of the temporal and occipital lobe. We argued that the target of the diffuse path is in a position to relate limbic and visual impulses and relay the product of such integration to the other visual areas, striate as well as extrastriate cortex.     

The projections from striate cortex (V1) to areas V2 and V3 in the macaque monkey: asymmetries, areal boundaries, and patchy connections

The organization of projections from V1 to areas V2 and V3 in the macaque monkey was studied with a combination of anatomical techniques, including lesions and tracer injections made in different portions of V1 and V2 in 20 experimental hemispheres. Our results indicate that dorsal V1 (representing the inferior contralateral visual quadrant) consistently projects in topographically organized fashion to V3 in the lunate and parietooccipital sulci as well as to the middle temporal area (MT) and dorsal V2. In contrast, ventral V1 (representing the superior contralateral quadrant) projects only to MT and ventral V2. A corresponding dorsoventral asymmetry in myeloarchitecture supports the idea that V3 is an area that is restricted to dorsal extrastriate cortex and lacks a complete representation of the visual field. The average surface area of myeloarchitectonically identified V3 was 89 mm2. Additional information was obtained concerning the laminar distribution of connections from V1 to V2 and V3, the patchiness of these projections, and the consistency of projections to other extrastriate areas, including V4 and V3A.     

Functional anatomy of the second visual area (V2) in the macaque

To study the functional organization of secondary visual cortex (V2) in the primate, 14C-2-deoxy-d-glucose (DG) was injected while macaque monkeys were shown specific visual stimuli. Wherever possible, patterns of DG uptake were compared with the position of dark and light cytochrome oxidase (cytox) stripes (Tootell et al., 1983). Often, the DG effects of 2 different stimuli were compared in the same hemisphere to eliminate ambiguities inherent in between-animal comparisons. Data were obtained from a large number of animals in conjunction with related DG studies in area V1 (primary visual or striate cortex). The following conclusions were reached: (1) in some macaque monkeys, dark cytox stripes were faint or absent. Although this could conceivably be due to poor staining technique, some evidence suggests that the lack of enzyme stripe pattern is real. In all animals, including those that showed poor or no cytox staining evidence for stripes, the functional architecture revealed by the DG was consistently present and robust. (2) Uniform gray stimuli produce a relatively uniform pattern and minimal stimulus-related DG uptake. (3) Eye movements per se produce some uptake in the V2 stripes. (4) Very generalized visual stimulation conditions (e.g., binocular stimulation with a grating of varied orientation and varied spatial frequency) produced a pattern of uptake that is greatest in both sets of dark cytox stripes and lighter in the light cytochrome stripes. (5) In both the DG and cytox results, the V2 "stripes" are more accurately described as stripe-shaped collections of patches. (6) In almost all cases, DG patterns were columnar in shape, extending from white matter to cortical surface. The boundaries of the columns were most sharply defined, and the contrast was highest, in layers 3B/4, becoming slightly more blurry and lower in contrast in other layers. Laminar differences between DG patterns in V2 were almost negligible, compared with the profound laminar differences in macaque V1. (7) There is no DG evidence for, and much against, the possibility of an ocular dominance architecture in V2. (8) There are orientation columns in macaque V2. DG-labeled orientation columns are spaced further apart than those in V1, by a factor of about 1.6, but the columns are not correspondingly wider. (9) Spatially diffuse variations in color produce high uptake confined, at least largely, to the thin cytox stripes. (10) There is evidence for spatially antagonistic color surrounds in color cells in the thin stripes.(ABSTRACT TRUNCATED AT 400 WORDS)     

Superior colliculus connections with visual thalamus in gray squirrels (Sciurus carolinensis): evidence for four subdivisions within the pulvinar complex

As diurnal rodents with a well-developed visual system, squirrels provide a useful comparison of visual system organization with other highly visual mammals such as tree shrews and primates. Here, we describe the projection pattern of gray squirrel superior colliculus (SC) with the large and well-differentiated pulvinar complex. Our anatomical results support the conclusion that the pulvinar complex of squirrels consists of four distinct nuclei. The caudal (C) nucleus, distinct in cytochrome oxidase (CO), acetylcholinesterase (AChE), and vesicular glutamate transporter-2 (VGluT2) preparations, received widespread projections from the ipsilateral SC, although a crude retinotopic organization was suggested. The caudal nucleus also received weaker projections from the contralateral SC. The caudal nucleus also projects back to the ipsilateral SC. Lateral (RLl) and medial (RLm) parts of the previously defined rostral lateral pulvinar (RL) were architectonically distinct, and each nucleus received its own retinotopic pattern of focused ipsilateral SC projections. The SC did not project to the rostral medial (RM) nucleus of the pulvinar. SC injections also revealed ipsilateral connections with the dorsal and ventral lateral geniculate nuclei, nuclei of the pretectum, and nucleus of the brachium of the inferior colliculus and bilateral connections with the parabigeminal nuclei. Comparisons with other rodents suggest that a variously named caudal nucleus, which relays visual inputs from the SC to temporal visual cortex, is common to all rodents and possibly most mammals. RM and RL divisions of the pulvinar complex also appear to have homologues in other rodents.