Retinotopic organization of striate and extrastriate visual cortex in the hooded rat

The visuotopic organization of the primary visual cortex (area 17) and the extrastriate visual regions surrounding it (areas 18a and 18) has been studied in gray rats using standard microelectrode mapping techniques. The results confirm and extend previous observations in the rat. Apart from the representation of the contralateral visual field (VF) in area 17, in which the upper VF is represented caudally and the nasal VF laterally, there are additional representations of the VF in the extrastriate cortex. In lateral extrastriate cortex (area 18a) there are at least 4 such representations, namely lateromedial (LM), anterolateral (AL), laterointermediate (LI) and laterolateral (LL). In LM (second visual area) the upper VF is represented caudally and the nasal VF medially, being thus a mirror image of V1. In AL (third visual area) the upper VF is represented rostrally and the nasal VF, medially, being thus a mirror image of LM. In LI, the upper VF is medial and the nasal VF, lateral, being thus a mirror image of LM, or a reduced copy of V1. In medial extrastriate cortex (area 18) there are two representations of the temporal VF, labeled anteromedial (AM) and posteromedial (PM). In AM, the upper temporal VF is medial and the lower temporal VF, lateral, the extreme temporal field being rostral. The 30° azimuth provides the boundary between AM and PM. Thus, AM is organized as a counter-clockwise rotation by 90° of the V1 representation. In PM, the upper lower VF topography is like in AM, but the extreme temporal VF is caudal, being thus a mirror image of AM.     

Resolving the organization of the New World monkey third visual complex: the dorsal extrastriate cortex of the marmoset (Callithrix jacchus)

We tested current hypotheses on the functional organization of the third visual complex, a particularly controversial region of the primate extrastriate cortex. In anatomical experiments, injections of retrograde tracers were placed in the dorsal cortex immediately rostral to the second visual area (V2) of New World monkeys (Callithrix jacchus), revealing the topography of interconnections between the "third tier" cortex and the primary visual area (V1). The data indicate the presence of a dorsomedial area (DM), which represents the entire upper and lower quadrants of the visual field, and which receives strong, topographically organized projections from the superficial layers of V1. The visuotopic organization and boundaries of DM were confirmed by electrophysiological recordings in the same animals and by architectural characteristics which were distinct from those found in ventral extrastriate cortex rostral to V2. There was no electrophysiological or histological evidence for a transitional area between V2 and DM. In particular, the central representation of the upper quadrant in DM was directly adjacent to the representation of the horizontal meridian that marks the rostral border of V2. The present results argue in favor of the hypothesis that the third visual complex in New World monkeys contains different areas in its dorsal and ventral components: area DM, near the dorsal midline, and a homolog of area 19 of other mammals, located more lateral and ventrally. The characteristics of DM suggest that it may correspond to visual area 6 (V6) of Old World monkeys.     

Relationships between interhemispheric cortical connections and visual areas in hooded rats

The correlation between visual topography within striate and lateral extrastriate visual cortex and the pattern of callosal connections to those areas has been studied in gray rats. The procedure was to put multiple injections of horseradish peroxidase (HRP) into the occipital cortex of the right hemisphere. The cortical areas 17 and 18a in the left hemisphere were electrophysiologically mapped upon stimulation of the right eye. Reference lesions were placed at selected recording sites. Horizontal sections of the left cortex were reacted for the demonstration of HRP. This permitted the comparison of the visual and callosal maps in the same animal. Like in other mammals, the callosal projections coincide with the cortical representations of the vertical midline and the more central regions of the visual field. The heavy line of labelled neurons and terminations embedded within the primary callosal band at the 17/18a border coincides with the representation of the vertical meridian. It provides the boundary between V1 and the maps located lateral to it. In area 18a, the anterolateral is contained in its anterior half, whereas areas lateromedial and laterointermediate are contained in its posterior half. The acallosal 'island', caudal to the acallosal 'body', contains the map known as posterolateral. There are two laterolateral (LL) maps which coincide with the acallosal 'islands' lateral to the acallosal 'body'; laterolateral anterior is more rostral than LL and these are retinotopically organized as mirror images of each other. Lateral to LL, there is a suggestion of an additional map, which could correspond to a pararhinal area. These results may be useful to understand aspects of a basic mammalian plan in the organization of visual cortex.     

Visual areas in lateral and ventral extrastriate cortices of the marmoset monkey

The representation of the visual field in visual areas of the dorsolateral, lateral, and ventral cortices was studied by means of extracellular recordings and fluorescent tracer injections in anaesthetised marmoset monkeys. Two areas, forming mirror-symmetrical representations of the contralateral visual field, were found rostral to the second visual area (V2). These were termed the ventrolateral posterior (VLP) and the ventrolateral anterior (VLA) areas. In both areas, the representation of the lower quadrant is located dorsally, between the foveal representation of V2 and the middle temporal crescent (MTc), whereas the representation of the upper quadrant is located ventrally, in the supratentorial cortex. A representation of the vertical meridian forms the common border of areas VLP and VLA, whereas the horizontal meridian is represented both at the caudal border of area VLP (with V2) and at the rostral border of area VLA (with multiple extrastriate areas). The foveal representations of areas VLP and VLA are continuous with that of V2, being located at the lateral edge of the hemisphere. The topographic and laminar patterns of projections from dorsolateral and ventral cortices to the primary (V1) and dorsomedial (DM) visual areas both support the present definition of the borders of areas VLP and VLA. These results argue against a separation between dorsolateral and ventral extrastriate areas and provide clues for the likely homologies between extrastriate areas of different species.     

Ventral posterior visual area of the macaque: visual topography and areal boundaries

Using both physiological and anatomical techniques, we have studied the topographic organization of extrastriate visual cortex on the ventral surface of the occipital lobe in macaque monkeys. Our results show that a topographically organized representation of the superior contralateral quadrant of the visual field lies immediately anterior to the ventral half of V2. This area is organized in a mirror symmetric fashion to ventral V2: it shares a horizontal meridian representation with V2 and a representation of the superior vertical meridian forms its anterior border. A well-defined strip of callosal inputs runs along the vertical meridian representation, thereby providing a reliable anatomical marker for areal boundaries in ventral extrastriate cortex. We refer to this area as the ventral posterior area (VP) because it is, in all these respects, notably similar to VP in the owl monkey. Ventral V2 has strong reciprocal connections with VP, and the topography of the V2 projection agrees closely with the topography revealed in our physiological mapping experiments. The visual field representation in VP is strikingly anisotropic, with linear magnification factor being much larger along contours of constant polar angle than along contours of constant eccentricity.     

Organization of visual cortex in the northern quoll, Dasyurus hallucatus: evidence for a homologue of the second visual area in marsupials

Two visual areas, V1 and V2 (first and second visual areas), appear to be present in the posterior neocortex of all eutherian mammals investigated so far. However, previous studies have not established whether an area homologous to V2 also exists in metatherian mammals (marsupials). Using electrophysiological techniques, we mapped the visual receptive fields of neurons in the striate and peristriate cortices of the northern quoll, an Australian marsupial. We found that neurons in a 2-mm-wide strip of cortex rostrolateral to V1 form a single, relatively simple representation of the complete contralateral hemifield. This area resembles V2 of eutherians in several respects: (i) neurons in the medial half of the peristriate area represent the lower visual quadrant, whereas those in the lateral half represent the upper visual quadrant; (ii) the vertical meridian of the visual field is represented adjacent to V1, while the visual field periphery is represented along the lateral and rostrolateral borders of the peristriate area; (iii) there is a marked anisotropy in the representation, with a larger magnification factor parallel to the V1 border than perpendicular to this border; and (iv) receptive fields of multiunit clusters in the peristriate cortex are much larger than those of cells in V1 at comparable eccentricities. The cortex immediately rostral and lateral to V2 did not respond to visual stimulation under our recording conditions. These results suggest that V1 and V2 together form a 'core' of homologous visual areas, likely to exist in all therian mammals.     

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.     

Central projections of the normal and ‘regenerate’ infraorbital nerve in adult rats subjected to neonatal unilateral infraorbital lesions: a transganglionic horseradish peroxidase study

The visuotopic organization of the primary visual cortex (area 17) and the extrastriate visual regions surrounding it (areas 18a and 18) has been studied in gray rats using standard microelectrode mapping techniques. The results confirm and extend previous observations in the rat. Apart from the representation of the contralateral visual field (VF) in area 17, in which the upper VF is represented caudally and the nasal VF laterally, there are additional representations of the VF in the extrastriate cortex. In lateral extrastriate cortex (area 18a) there are at least 4 such representations, namely lateromedial (LM), anterolateral (AL), laterointermediate (LI) and laterolateral (LL). In LM (second visual area) the upper VF is represented caudally and the nasal VF medially, being thus a mirror image of V1. In AL (third visual area) the upper VF is represented rostrally and the nasal VF, medially, being thus a mirror image of LM. In LI, the upper VF is medial and the nasal VF, lateral, being thus a mirror image of LM, or a reduced copy of V1. In medial extrastriate cortex (area 18) there are two representations of the temporal VF, labeled anteromedial (AM) and posteromedial (PM). In AM, the upper temporal VF is medial and the lower temporal VF, lateral, the extreme temporal field being rostral. The 30 degrees azimuth provides the boundary between AM and PM. Thus, AM is organized as a counter-clockwise rotation by 90 degrees of the V1 representation. In PM, the upper lower VF topography is like in AM, but the extreme temporal VF is caudal, being thus a mirror image of AM.     

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

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

A systematic topographical relationship between mouse lateral posterior thalamic neurons and their visual cortical projection targets

Higher-order visual thalamus communicates broadly and bi-directionally with primary and extrastriate cortical areas in various mammals. In primates, the pulvinar is a topographically and functionally organized thalamic nucleus that is largely dedicated to visual processing. Still, a more granular connectivity map is needed to understand the role of thalamocortical loops in visually guided behavior. Similarly, the secondary visual thalamic nucleus in mice (the lateral posterior nucleus, LP) has extensive connections with cortex. To resolve the precise connectivity of these circuits, we first mapped mouse visual cortical areas using intrinsic signal optical imaging and then injected fluorescently tagged retrograde tracers (cholera toxin subunit B) into retinotopically-matched locations in various combinations of seven different visual areas. We find that LP neurons representing matched regions in visual space but projecting to different extrastriate areas are found in different topographically organized zones, with few double-labeled cells (~4–6%). In addition, V1 and extrastriate visual areas received input from the ventrolateral part of the laterodorsal nucleus of the thalamus (LDVL). These observations indicate that the thalamus provides topographically organized circuits to each mouse visual area and raise new questions about the contributions from LP and LDVL to cortical activity.     

The evolution of visual cortex: where is V2?

A comparative analysis of the area of the cortex that is adjacent to the primary visual area (V1), indicates that the lateral extrastriate cortex of primitive mammals was likely to contain only a single visuotopically organized field, the second visual area (V2). Few, if any, other visual areas existed. The opposing hypothesis, that primitive mammals had a 'string' of small visual areas in the cortex lateral to V1 (as in some rodents), is not supported by studies of the organization of extrastriate cortex in other mammals, nor by the variability in this organization among extant rodents. A critical re-analysis of published evidence on the presence of multiple areas adjacent to V1 in some rodents has led to alternative interpretations of the organization of the areas in these regions.     

Retinal lesions induce fast intrinsic cortical plasticity in adult mouse visual system

Neuronal activity plays an important role in the development and structural–functional maintenance of the brain as well as in its life‐long plastic response to changes in sensory stimulation. We characterized the impact of unilateral 15° laser lesions in the temporal lower visual field of the retina, on visually driven neuronal activity in the afferent visual pathway of adult mice using in situ hybridization for the activity reporter gene zif268. In the first days post‐lesion, we detected a discrete zone of reduced zif268 expression in the contralateral hemisphere, spanning the border between the monocular segment of the primary visual cortex (V1) with extrastriate visual area V2M. We could not detect a clear lesion projection zone (LPZ) in areas lateral to V1 whereas medial to V2M, agranular and granular retrosplenial cortex showed decreased zif268 levels over their full extent. All affected areas displayed a return to normal zif268 levels, and this was faster in higher order visual areas than in V1. The lesion did, however, induce a permanent LPZ in the retinorecipient layers of the superior colliculus. We identified a retinotopy‐based intrinsic capacity of adult mouse visual cortex to recover from restricted vision loss, with recovery speed reflecting the areal cortical magnification factor. Our observations predict incomplete visual field representations for areas lateral to V1 vs. lack of retinotopic organization for areas medial to V2M. The validation of this mouse model paves the way for future interrogations of cortical region‐ and cell‐type‐specific contributions to functional recovery, up to microcircuit level.     

Topographic organization of human visual areas in the absence of input from primary cortex.

Recently, there has been evidence for considerable plasticity in primary sensory areas of adult cortex. In this study, we asked to what extent topographical maps in human extrastriate areas reorganize after damage to a portion of primary visual (striate) cortex, V1. Functional magnetic resonance imaging signals were measured in a subject (G.Y.) with a large calcarine lesion that includes most of primary visual cortex but spares the foveal representation. When foveal stimulation was present, intact cortex in the lesioned occipital lobe exhibited conventional retinotopic organization. Several visual areas could be identified (V1, V2, V3, V3 accessory, and V4 ventral). However, when stimuli were restricted to the blind portion of the visual field, responses were found primarily in dorsal extrastriate areas. Furthermore, cortex that had formerly shown normal topography now represented only the visual field around the lower vertical meridian. Several possible sources for this reorganized activity are considered, including transcallosal connections, direct subcortical projections to extrastriate cortex, and residual inputs from V1 near the margin of the lesion. A scheme is described to explain how the reorganized signals could occur based on changes in the local neural connections.     

Interhemispheric connections of visual cortex in the owl monkey,Aotus trivirgatus,and the bushbaby,Galago senegalensis

Anatomical techniques have been used to map within visual cortex the pattern of degenerating axonal terminals produced by surgical section of the splenium of the corpus callosum in the owl monkey, Aotus trivirgatus, and the bushbaby, Galago senegalensis. Previous studies in other species have shown that callosal inputs terminate preferentially in regions where the vertical meridian of the visual field is represented. Such a correspondence can serve as a useful aid for locating the boundaries of visual areas. The goals of this study have been (1) to assess the degree of correspondence between callosal inputs and previously identified vertical meridian representations in the owl monkey and bushbaby, and (2) to gain information from the pattern of callosal inputs concerning the existence and organization of as yet unidentified extrastriate visual areas. In both the owl monkey and the bushbaby, a discrete band of degenerating axonal terminals corresponds precisely to the vertical meridian representation at the V1-V2 border, and a less precise increase in the density of degenerating axonal terminals corresponds to the vertical meridian representation of extrastriate area MT. A well-defined band of degeneration on the ventral surface of the owl monkey's cerebral hemisphere corresponds to a previously unknown vertical meridian representation which is shared by two newly identified extrastriate visual areas. Elsewhere in visual cortex the pattern of callosal connections is more complex. Although this pattern may still reflect visual topography, it is not immediately useful for distinguishing areal boundaries.     

Topographical organization of cortical afferents to extrastriate visual area PO in the macaque: a dual tracer study

We have examined the origin and topography of cortical projections to area PO, an extrastriate visual area located in the parieto-occipital sulcus of the macaque. Distinguishable retrograde fluorescent tracers were injected into area PO at separate retinotopic loci identified by single-neuron recording. The results indicate that area PO receives retinotopically organized inputs from visual areas V1, V2, V3, V4, and MT. In each of these areas the projection to PO arises from the representation of the periphery of the visual field. This finding is consistent with neurophysiological data indicating that the representation of the periphery is emphasized in PO. Additional projections arise from area MST, the frontal eye fields, and several divisions of parietal cortex, including four zones within the intraparietal sulcus and a region on the medial dorsal surface of the hemisphere (MDP). On the basis of the laminar distribution of labeled cells we conclude that area PO receives an ascending input from V1, V2, and V3 and receives descending or lateral inputs from all other areas. Thus, area PO is at approximately the same level in the hierarchy of visual areas as areas V4 and MT. Area PO is connected both directly and indirectly, via MT and MST, to parietal cortex. Within parietal cortex, area PO is linked to particular regions of the intraparietal sulcus including VIP and LIP and two newly recognized zones termed here MIP and PIP. The wealth of connections with parietal cortex suggests that area PO provides a relatively direct route over which information concerning the visual field periphery can be transmitted from striate and prestriate cortex to parietal cortex. In contrast, area PO has few links with areas projecting to inferior temporal cortex. The pattern of connections revealed in this study is consistent with the view that area PO is primarily involved in visuospatial functioning.     

Cortical projections of the dorsolateral visual area in owl monkeys: the prestriate relay to inferior temporal cortex

The dorsolateral visual area (DL) is one of a number of visual areas that have been defined by electrophysiological mapping procedures and cortical architecture in the extrastriate cortex of owl monkeys. The projections of DL were determined by the intra-axonal transport of 3H-proline, 3H-acetyl-wheat germ agglutinin, and horseradish peroxidase after cortical injections. The major ipsilateral projection of DL defined a new subdivision of the visual cortex in owl monkeys, the caudal inferior temporal cortex. Single injections in DL sometimes produced label in two separate regions in the caudal inferior temporal cortex, suggesting that functional subdivisions exist in this projection zone. Other targets of DL included the region of the frontal eye fields, the dorsomedial visual area, the dorsointermediate visual area (DI), a region of the cortex rostral to DI which we call the temporoparietal cortex, and possibly the ventral (V) and posterior parietal areas. A major feedback projection of DL was to V-II. Projections from DL to V-II and the dorsomedial visual area were roughly retinotopic. Projections from DL to the contralateral cerebral hemisphere were to DL and the inferior temporal cortex. Overall, the results support the concept that a major relay of visual information proceeds from V-I to V-II to DL and then to the inferior temporal cortex. In addition, similarities in connection patterns of DL in owl monkeys and V4 in macaque monkeys suggest that DL and much or all of V4 are homologous.     

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.     

Occipital cortex in man: organization of callosal connections, related myelo- and cytoarchitecture, and putative boundaries of functional visual areas

Human area 17 is known to contain a single (the primary) visual area, whereas areas 18 and 19 are believed to contain multiple visual areas (defined as individual representations of the contralateral visual hemifield). This is known to be the case in monkeys, where several boundaries between visual areas are characterized by bands of callosal afferents and/or by changes in myeloarchitecture. We here describe the pattern of callosal afferents in (human) areas 17, 18, and 19 as well as their cortical architecture and we infer the position of some visual areas. Sections from occipital lobes of 6 human brains with unilateral occipital infarctions have been silver-impregnated for degenerating axons, thereby revealing callosal afferents to the intact occipital cortex. Their tangential distribution is discontinuous, even in cases with large lesions. A band of callosal afferents straddles the area 17/18 boundary, whereas the remainder of area 17 and a 15-45 mm wide stripe of area 18 adjacent to the callosal band along the 17/18 border are free of them. Patches of callosal afferents alternate with callosal-free regions more laterally in area 18 and in area 19. We conclude that, in man, a second visual area (analogue of V2) lies in area 18, horseshoe-shaped around area 17, and includes the inner part of the acallosal stripe adjacent to the callosal band along the 17/18 boundary. The outer part of this acallosal stripe belongs to a third visual area, which may contain dorsally the analogue of V3 and ventrally that of VP. Thus the lower parts of the second and third visual areas lie on the lingual gyrus, whereas the analogue of the macaque's fourth visual area probably lies on the fusiform gyrus. Although the proposed subdivision of the occipital cortex relies largely on the pattern of callosal afferents, some putative human visual areas appear to have distinct architectonic features. The analogue of V2 is rather heavily myelinated and its layer III contains large pyramidal neurons. Its upper part is not well delimited laterally since adjacent "V" has similar architecture. Its lower part, however, differs clearly from the adjacent "VP," which is lightly myelinated and lacks the large pyramids in layer III. The cortex lateral to "VP" is heavily myelinated and contains fairly large pyramids in layers III and V. The myeloarchitecture of the lateral part of the occipital cortex is not uniform; a very heavily myelinated region stands out in the lateral part of area 19, near the occipito-temporal junction.(ABSTRACT TRUNCATED AT 400 WORDS)     

Functional Differentiation of Mouse Visual Cortical Areas Depends upon Early Binocular Experience

The mammalian visual cortex contains multiple retinotopically defined areas that process distinct features of the visual scene. Little is known about what guides the functional differentiation of visual cortical areas during development. Recent studies in mice have revealed that visual input from the two eyes provides spatiotemporally distinct signals to primary visual cortex (V1), such that contralateral eye-dominated V1 neurons respond to higher spatial frequencies than ipsilateral eye-dominated neurons. To test whether binocular visual input drives the differentiation of visual cortical areas, we used two-photon calcium imaging to characterize the effects of juvenile monocular deprivation (MD) on the responses of neurons in V1 and two higher visual areas, LM (lateromedial) and PM (posteromedial). In adult mice of either sex, we find that MD prevents the emergence of distinct spatiotemporal tuning in V1, LM, and PM. We also find that, within each of these areas, MD reorganizes the distinct spatiotemporal tuning properties driven by the two eyes. Moreover, we find a relationship between speed tuning and ocular dominance in all three areas that MD preferentially disrupts in V1, but not in LM or PM. Together, these results reveal that balanced binocular vision during development is essential for driving the functional differentiation of visual cortical areas. The higher visual areas of mouse visual cortex may provide a useful platform for investigating the experience-dependent mechanisms that set up the specialized processing within neocortical areas during postnatal development.SIGNIFICANCE STATEMENT Little is known about the factors guiding the emergence of functionally distinct areas in the brain. Using in vivo Ca²⁺ imaging, we recorded visually evoked activity from cells in V1 and higher visual areas LM (lateromedial) and PM (posteromedial) of mice. Neurons in these areas normally display distinct spatiotemporal tuning properties. We found that depriving one eye of normal input during development prevents the functional differentiation of visual areas. Deprivation did not disrupt the degree of speed tuning, a property thought to emerge in higher visual areas. Thus, some properties of visual cortical neurons are shaped by binocular experience, while others are resistant. Our study uncovers the fundamental role of binocular experience in the formation of distinct areas in visual cortex.     

Retinotopic mapping reveals extrastriate cortical basis of homonymous quadrantanopia

It has been conventionally assumed that cortically based quadrantic visual field deficits (homonymous quadrantanopias) are caused by lesions in striate cortex (V1), extending precisely to the horizontal meridian representation. A more recent model, supported by anatomic MRI evidence, consists of an exclusively extrastriate cortical basis (e.g. V2, V3, VP, V4v). Employing fMRI, we sought to distinguish between these models through retinotopic mapping of a patient with an upper right homonymous quadrantanopia. As expected, maps of the lower right quadrant and left hemifield were normal. The map corresponding to the impaired upper right quadrant was normal in V1 and V2, with little or no activity in VP and V4v. These results provide functional evidence that extrastriate cortical lesions can elicit homonymous quadrantanopias.