Functional Specificity of Callosal Connections in Tree Shrew Striate Cortex

Although callosal connections have been shown to link extensive regions of primary visual cortex, the distribution of these connections with respect to the map of visual space and the map of orientation preference remains unclear. Here we combine optical imaging of intrinsic signals with injection of fluorescent microspheres to assess the functional specificity of callosal connections in the tree shrew. By imaging both hemispheres simultaneously while presenting a series of spatially restricted stimuli, we find that a substantial region of visual space is represented bilaterally. Each hemisphere includes a representation of the ipsilateral visual field that is highly compressed relative to that of the contralateral visual field and is most extensive in the lower visual field, where ∼30ο of central visual space are represented bilaterally. Callosal connections extend throughout the region of bilateral representation but terminate in a spatially restricted manner that links visuotopically corresponding sites in the two hemispheres. In contrast, callosal connections appear to terminate without regard for the map of orientation preference, showing little sign of the orientation-specific modular and axial specificity that is characteristic of long-range horizontal connections. By coordinating the activity in the two hemispheres in a way that preserves nearest neighbor relationships, callosal connections may best be viewed as elements of local circuits that operate within a single bilateral representation of visual space.

Related papers:

title: THE FUNCTION OF THE CALLOSAL CONNECTIONS OF THE VISUAL CORTEX.
abstract: There is anatomical evidence in cats and baboons of cells at the margin of area 17 which have callosal connections to corresponding points in the opposite hemisphere. The function of such a pathway has been studied in cats after one optic tract has been cut. The corresponding hemisphere now gives early visual responses only at the margin of area 17. Such responses can only be elicited by light stimuli near the vertical meridian of the visual field. These responses are abolished by cooling the corresponding points in the opposite hemisphere and by cutting the corpus callosum. This pathway appears to provide some functional union for the two halves of the visual field.

title: Cortical and callosal connections concerned with the vertical meridian of visual fields in the cat.
abstract: IN HIGHER MAMMALS the left half of the visual field is represented in the right visual cortex, and the right half in the left cortex. The fact that our visual fields appear uniform, with no obvious interruption along the vertical midline, would seem to call for connections linking the two hemispheres. The present paper deals with the physiological properties of some of these connections. In the cat the vertical midline of the visual field has its primary representation in and close to the poorly defined boundary between areas 17 and 18 (7, 10). Each half-visual field is represented in both 17 and 18 in mirror fashion, so that as one proceeds away from the 17-18 boundary in either direction, medially along 17 or laterally along 18, the corresponding region of visual fields moves outward from the vertical meridian into the contralateral visual field. Recent anatomical evidence indicates that the part of area 17 adjacent to the 17-18 boundary sends callosal projections to the other hemisphere, ending in 18 near the 17-18 boundary (2,7,8) and possibly also in the adjacent part of 17 (8). Thus the parts of 17 and 18 having to do with the vertical midline of the visual fields are apparently connected on the two sides by the corpus callosum. In contrast, the more medial parts of 17, representing the peripheral parts of the visual fields, seem not to be reciprocally interconnected, and indeed one would hardly expect that cells concerned with a particular part of the left visual field should be connected specifically to cells concerned with the corresponding region in the right field. These anatomical findings have lately been confirmed in the elegant physiological experiments of Choudhury, Whitteridge, and Wilson (1). They showed that with one opti c trac t cut responses to light could be evoked in the ipsilateral hemisphere only near the 17-18 border, and only when the stimulus was applied near the vertical meridian. The responses were abolished either by cooling the corresponding region in the contralateral hemisphere or by cooling or cutting the posterior corpus callosum. Several lines of evidence thus indicate that fibers in the corpus callosum originating in the most lateral part of 17 and projecting to 18 on the other side have special functions involving the vertical midline of the visual field.

title: Binocularly Driven Neurons in Visual Cortex of Split-Chiasm Cats
abstract: In cats with midsagittal section of the optic chiasm, some visual cortex neurons can be driven not only by the ipsilateral eye, through the direct geniculocortical pathways, but also by the contralateral eye, through the opposite visual cortex and corpus callosum. The receptive fields and the response characteristics observed upon stimulation of the contralateral eye are very similar to those observed upon stimulation of the ipsilateral eye; the two monocular receptive fields of a given cell lie in corresponding points of heteronymous halves of the visual field in close contact with the vertical meridian, thus adding in visual space and forming a binocular receptive area which crosses the vertical meridian and extends equally on either side of it.

title: The cells of origin of the corpus callosum in rabbit visual cortex
abstract: It has become apparent in recent years that there are significant species differences in both the laminar and regional distribution of the cell bodies o f callosal efferent neurons (CENs) which originate in visual cortex (see e.g. refs. 3,4,6,9,11,13 and 14). In the present study we combine electrophysiological mapping procedures with the use of horseradish peroxidase (HRP) in order to examine the regional and laminar distribution of CENs in visual areas I and II (V-I and V-II, respectively) of rabbit. In this species the border between physiologically defined V-I and V-II zz may correspond to the border between cytoarchitectonically defined striate and occipital cortex v (see discussion by Towns et al.la). This correspondence, however, has not yet been established. Consequently, we physiologically defined the border between V-I and V-II in most of the cases described below. Successful cases utilized 17 adult (6-24-month) Dutch rabbits. In the first experimental paradigm , multiple injections of H R P were made th roughout much of

title: Interhemispheric connections of cortical sensory areas in tree shrews
abstract: Interhemispheric connections were studied in tree shrews (Tupaia belangeri) after multiple injections of horseradish peroxidase or horseradish peroxidase conjugated to wheat germ agglutinin into the cortex of one cerebral hemisphere. After an appropriate survival period, the areal pattern of connections was revealed by flattening the other hemisphere, cutting sections parallel to the cortical surface, and staining with tetramethylbenzidine. Architectonic boundaries were identified by using sections stained for myelinated fibers. Labeled cells and axon terminations formed largely overlapping distributions that covaried in density, although labeled cells appeared to be more evenly distributed than labeled terminations. Connections were concentrated along the border of area 17 (V‐I) with area 18 (V‐II). However, connections also extended as far as 2 mm into area 17 to include cortex representing parts of the visual field 10° or more from the zero vertical meridian. Clusters of dense connections spanned the width of area 18, where they alternated with regions of fewer connections. These clusters roughly corresponded in location to regions with heavier myelination. In the visually responsive temporal cortex, connections were also unevenly distributed. The organization of most of this cortex is not understood, but one subdivision, the temporal dorsal area (TD), has been identified on the basis of reciprocal connections with area 17. The central part of the TD had few interhemispheric connections, while most of the outer border had dense connections. The auditory cortex had dense and patchy connections throughout. The pattern in the primary somatosensory cortex (S‐I) varied according to the representation of body parts, so that the cortex related to the forepaw had sparse connections, while connections were dense but uneven over much of the representation of the face, nose, and mouth. A focus of connections was found at the border of the forepaw and face representations, where the myelination of S‐I cortex is interrupted. Dense, uneven connections also characterized the second somatosensory area, S‐II. The motor cortex was densely connected, with only slightly fewer terminations rostral to the forepaw region of S‐I. Other parts of frontal cortex had dense connections, The distribution of cortical connections varied with depth for at least some areas, so that clusters of cells and terminations were found in supragranular layers in S‐I, S‐II, and TD, while infragranular labeled cells were more evenly distributed.

title: Cortical connections of area 17 in tree shrews
abstract: In order to better understand the organization of extrastriate cortex in tree shrews, injections in area 17 of wheat germ agglutinin or tritiated proline were used to reveal an intrinsic pattern of connections, ipsilateral connections with area 18 and two other subdivisions of cortex, and callosal connections with areas 17 and 18 of the opposite cerebral hemisphere. Areal patterns of connections were best seen in sections cut parallel to the surface of flattened cortex. Within area 17, periodic foci of labeled terminations and cells extended from and surrounded injection sites as described by Rockland et al. ('82). Single injections produced multiple foci of labeled terminations and cells in area 18. The foci tended to fuse into short bands that sometimes crossed the width of area 18. Double injections produced more foci, and multiple injections tended to produce more continuous regions of label. An overall retinotopic pattern was evident with rostral area 17 connected to rostral area 18 and caudal area 17 connected to caudal area 18. Terminations extended through layers II–VI, with some increase in density in layer IV. Cells in area 18 projecting back to area 17 were in layers III and V. The injections also allowed identification of previously undefined subdivisions of visual cortex in temporal cortex immediately adjoining area 18. Dense reciprocal connections were observed in a 13 mm2 oval of cortex on the lateral border of the middle section of area 18 that we define as the temporal dorsal area, TD. Connections indicate a crude topographic organization with lower field represented rostrally and upper field caudally. Inputs were most dense in the middle cortical layers, and labeled cells were supragranular, and less frequently, infragranular. A 10‐mm2 oval of cortex near the posterior edge of the hemisphere, the temporal posterior area (TP), contained labeled cells after area 17 injections, but terminal labeling was only obvious in the dorsal part. Single injections sometimes produced quite separate dorsal and ventral zones of label in TP, suggesting a small separate dorsal division. A crude retinotopic order appears to exist within ventral TP, with the lower field most ventral. Labeled cells were largely supragranular. A fourth zone of ipsilateral connections was in posterior limbic cortex bordering area 17 on the ventromedial surface of the cerebral hemisphere. The callosal connections were reciprocal and included regions 1 mm wide on either side of the area 17 and area 18 border. Callosal connections were roughly homotopic. Callosal terminations included superficial layers, and projecting cells were both supragranular and infragranular. We conclude that extrastriate cortex in tree shrews contains at least three subdivisions, one of which (TD) may be the homologue of the middle temporal visual area of primates.

title: Callosal projections between areas 17 in the adult tree shrew (Tupaia belangeri)
abstract: SummaryIn the primary visual cortex (area 17) of the tree shrew (Tupaia belangeri) neurons projecting to the contralateral area 17 via the corpus callosum were identified by horseradish peroxidase histochemistry (HRP, WGA-HRP). The distribution of homotopic and heterotopic connections was studied. We found that a narrow stripe of area 17 close to the dorsal area 17/18 border — which corresponds to the visual field along the vertical meridian — is connected via homotopic callosal projections. The adjacent dorsal part of area 17, which largely corresponds to the binocular visual field, is connected via homotopic as well as heterotopic projections. Heterotopic projections originate in the cortical stripe along the area 17/18 border and their contralateral targets are displaced medially. Callosal neurons are located mostly in supragranular but also occur in infragranular layers. The supragranular neurons in general are pyramidal cells. In addition to these findings, we confirmed earlier reports on ipsilateral projections of the primary visual area to the dLGN, the claustrum, area 18 and other visual areas.

title: Reciprocal heterotopic callosal connections between the two striate areas in Tupaia
abstract: SummaryWGA-HRP injections were placed into area 17 close to the border with area 18 of Tupaia belangeri in order to study the callosal connections of the striate area in this animal. Most callosal neurons were found in the striate cortex (57.6–86.9%), some in the extrastriate area 18 (10.6–28.1%), and a few in even more temporal regions (2.5–14.3%). Concerning only the area 17, reciprocal homotopic connections could be observed as a strip along the area 17/18 border. Additionally, heterotopic callosal connections could be seen in regions representing the binocular visual field, especially the lower part. The area 17 cells were mostly located in the supragranular layers II and III (94.1–97.2%). But neurons could also be found in the infragranular layers, especially layer VI (2.6–5.2%) and in layer IV (0.2–1.1%). Homotopic projections were mostly seen in layers IIIc and V. The majority of the supragranular and infragranular neurons are pyramidal cells. However, a newly defined subpopulation of neurons, most probably stellate cells, were discovered forming a band in sublayer IIIc, very close to the layer III/IV border.

title: Visual-field map in the transcallosal sending zone of area 17 in the cat.
abstract: The representation of the visual field in the part of area 17 containing neurons that project axons across the corpus callosum to the contralateral hemisphere was defined in the cat. Of 1424 sites sampled along 77 electrode tracks, 768 proved to be in the callosal sending zone, which was identified by retrograde transport of horseradish peroxidase that had been deposited in the opposite hemisphere. The results show that the callosal sending zone has a fairly constant width of between 3 and 4 mm at most levels in area 17. However, the representation of the contralateral field at the different elevations of the visual field is not equal in this zone. The zone represents positions within 4 deg of the midline at the 0-deg horizontal meridian, and positions out to 15-deg azimuths in the upper hemifield and out to positions of 25-deg azimuth in the lower hemifield. The shape of the representation is approximately mirror-symmetric about the horizontal meridian, although there is a greater extent in the lower hemifield, which can be accounted for by the greater range of elevations (greater than 60 deg) represented there compared with the upper hemifield (approximately 40 deg). The representation in the sending zone of one hemisphere matches that present in the area 17/18 transition zone, which receives the bulk of transcallosal projections, in the opposite hemisphere. The observations on the sending zone show that callosal connections of area 17 are concerned with a vertical hour-glass-shaped region of the visual field centered on the midline. The observations suggest that in addition to interactions between neurons concerned with positions immediately adjacent to the midline, there are positions, especially high and low in the visual field, where interactions can occur between neurons that have receptive fields displaced some distance from the midline.

title: Visual-field map in the callosal recipient zone at the border between areas 17 and 18 in the cat.
abstract: The representation of the visual field in the callosal fiber recipient zone of area 17 and the adjacent area 17/18 transition zone was determined in the cat. The callosal fiber recipient zone was identified by anterograde transport of tritiated amino acids that had been injected into transcallosal sending zone of the opposite hemisphere. Application of autoradiographic procedures revealed that transcallosal projections are densest in the area 17/18 transition zone, and that their density in area 17 diminishes within 1-2 mm of the transition zone. Of 980 sites sampled in the visual-field mapping part of the study, 507 proved to be in the zone demarcated by transcallosally transported label. In this zone, both ipsilateral- and contralateral-field positions are represented, and the representation of the visual field at the different elevations is not equal. When ipsilateral-field positions are considered, the representation extends to about 4 deg close to the visual axis, and to 15-20 deg at elevations greater than +/- 30 deg, the representation is approximately mirror-symmetric about the horizontal meridian, and the representation is concordant with that of the representation in the area 17 transcallosal sending zone of the opposite hemisphere.

title: Visual callosal projections in the adult ferret.
abstract: The laminar and tangential organization of visual callosal projections of areas 17 and 18 were investigated in the adult ferret, using histochemical methods to visualize axonally transported horseradish peroxidase (HRP). Normal adult ferrets were given injections of HRP throughout one visual cortex or had gelfoam soaked in HRP applied to the transected corpus callosum. The ferret callosal cell distribution has a greater tangential extent in area 18 than in area 17. In addition, the radial organization of callosal cells in areas 17 and 18 differs: three times as many infragranular cells are present in area 18 than in area 17, although the number of supragranular cells is similar for both areas 17 and 18. Since the projections of alpha retinal ganglion cells are reported to be exclusively contralateral in the ferret (Vitek et al., 1985), callosal projections may make a major contribution to the binocularity of neurons in area 18.

title: Widespread Periodic Intrinsic Connections in the Tree Shrew Visual Cortex
abstract: Intrinsic connections within the tree shrew (Tupaia glis) visual cortex (area 17) are organized in periodic stripelike patterns within layers I, II, and III. This anatomical network resembles the regularly organized stripes of 2-deoxyglucose accumulation seen after stimulation of alert animals with uniformly oriented lines. Such connections imply that widespread lateral interactions are superimposed on the retinotopic organization ofarea 17 and suggest alternative interpretations ofcortical 1532 on July 1, 2017 http://sce.sciencem agorg/ D ow nladed fom

title: General Organization of Callosal Connections in the Cerebral Cortex
abstract: The necessity of interhemispheric connections, and the nature of this necessity, are demonstrated by the following hypothetical event. An intelligent being from outer space lands on earth and is asked to design the brain of a cat. The being is intrigued to find that the body of a cat is bilaterally symmetric (he looks himself rather like a multieyed and multiwhiskered octopus).

title: Organization of the callosal connections of visual areas v1 and v2 in the macaque monkey
abstract: The interhemispheric efferent and afferent connections of the V1/V2 border have been examined in the adult macaque monkey with the tracers horseradish peroxidase and horseradish peroxidase conjugated to wheat germ agglutinin. The V1/V2 border was found to have reciprocal connections with the contralateral visual area V1, as well as with three other cortical sites situated in the posterior bank of the lunate sulcus, the anterior bank of the lunate sulcus, and the posterior bank of the superior temporal sulcus.

title: Morphology of Callosal Axons Interconnecting Areas 17 and 18 of the Cat
abstract: Seventeen callosally projecting axons originating near the border between areas 17 and 18 in adult cats were anterogradely labelled with biocytin and reconstructed in 3‐D from serial sections. All axons terminated near the contralateral 17/18 border. However, they differed in their diameter, tangential and radial distributions, and overall geometry of terminal arbors. Diameters of reconstructed axons ranged between 0.45 and 2.25 μm. Most of the axons terminated in multiple terminal columns scattered over several square millimetres of cortex. Thus in general callosal connections are not organized according to simple, point‐to‐point spatial mapping rules. Usually terminal boutons were more numerous in supragranular layers; some were also found in infragranular layers, none in layer IV. However, a few axons were distributed only or mainly in layer IV, others included this layer in their termination. Thus, different callosal axons may selectively activate distinct cell populations. The geometry of terminal arbors defined two types of architecture, which were sometimes represented in the same axon: parallel architecture was characterized by branches of considerable length which supplied different columns or converged onto the same column; serial architecture was characterized by a tangentially running trunk or main branch with radial collaterals to the cortex. These architectures may relate to temporal aspects of inter‐hemispheric interactions. In conclusion, communication between corresponding areas of the two hemispheres appears to use channels with different morphological and probably functional properties.

title: Functional Specificity of Long-Range Intrinsic and Interhemispheric Connections in the Visual Cortex of Strabismic Cats
abstract: The development of both long-range intracortical and interhemispheric connections depends on visual experience. Previous experiments showed that in strabismic but not in normal cats, clustered horizontal axon projections preferentially connect cell groups activated by the same eye. This indicates that there is selective stabilization of fibers between neurons exhibiting correlated activity. Extending these experiments, we investigated in strabismic cats: (1) whether tangential connections remain confined to columns of similar orientation preference within the subsystems of left and right eye domains; and (2) whether callosal connections also extend predominantly between neurons activated by the same eye and preferring similar orientations. To this end, we analyzed in strabismic cats the topographic relationships between orientation preference domains and both intrinsic and callosal connections of area 17. Red and green latex microspheres were injected into monocular iso-orientation domains identified by optical imaging of intrinsic signals. Additionally, domains sharing the ocular dominance and orientation preference of the neurons at the injection sites were visualized by 2-deoxyglucose (2-DG) autoradiography. Quantitative analysis revealed that 56% of the retrogradely labeled cells within the injected area 17 and 60% of the transcallosally labeled neurons were located in the 2-DG-labeled iso-orientation domains. This indicates: (1) that strabismus does not interfere with the tendency of long-range horizontal fibers to link predominantly neurons of similar orientation preference; and (2) that the selection mechanisms for the stabilization of callosal connections are similar to those that are responsible for the specification of the tangential intrinsic connections.

title: Non‐mirror‐symmetric patterns of callosal linkages in areas 17 and 18 in cat visual cortex
abstract: In the cat, callosal connections in area 17 are largely confined to a 5–6‐mm‐wide strip at the 17/18 border. It is commonly thought that callosal fibers extending from between the 17/18 border regions interconnect loci that are mirror‐symmetric with respect to the midline of the brain, but this idea has not been tested experimentally. The present study examined the organization of callosal linkages in the 17/18 border region of normal adult cats by analyzing the patterns of connections revealed in one hemisphere after small injections of different fluorescent tracers into the opposite 17/18 callosal region. The location of the injection sites within areas 17 and 18 was assessed by examining architectonic data and by inspecting the labeling pattern in the ipsilateral visual thalamus. Area 17 and 18 were separated by a 1 –1.5‐mm‐wide zone of cytoarchitectonic transition rather than by a sharp border.

title: The organization of binocular cortex in the primary visual area of the rabbit
abstract: A conscious rabbit which crouches in the ‘freeze’ position has an unequivocal 24° wide binocular field formed by the overlap of the two 12° sectors of uniocular optical field which extend nasal of its midline. Although this investigation reveals that the horizontal representation of the uniocular visual field in cortical visual area I extends more nasal than in earlier reports, it is found to terminate at the midline when the eyes are set in the standard freeze position. The 12° sectors of uniocular field nasal of the midline were not represented in spite of being served by retina. The binocular field of the rabbit in the freeze posture thus appears to have no binocular representation in visual area I.

title: Ipsilateral visual field represented in the cat's visual cortex
abstract: Responses to visual stimulation were recorded in cells from the border region between cortical areas 17 and 18 in anaesthetized cats. There was found to be a band of ipsilateral representation, the cells of which had rather large receptive fields centered up to 8-12 degrees into the ipsilateral hemifield. The edges of the receptive fields extended 2-3 degrees still further ipsilaterally. The cortical region involved is very small, however, stretching 0.2-1.0 mm mediolaterally, and possibly restricted largely to layers II and III. Hence, in cats, as in sheep and hamsters, there is a band about 20 degrees wide of visual field along the naso-temporal division that is represented in the visual cortices of both hemispheres.

title: Maps of Central Visual Space in Ferret V1 and V2 Lack Matching Inputs from the Two Eyes
abstract: In the visual cortex, the representation of central visual space is supplied by matching geniculate inputs that are driven exclusively by one eye or the other. In layer 4 of early visual areas (V1 in primates and V1 and V2 in cat), these inputs form a nearly uniform array of small ocular dominance domains, while preserving overall topographic order within the cortical map. In ferret, however, ocular dominance domains in different regions of the visual cortex are strikingly irregular in size and shape. The exceptionally large size of domains in some regions implies a departure from the usual visuotopic matching of inputs from the two eyes. Using optical-imaging, electrophysiological, and anatomical techniques, we show that this regional variation is attributable to exclusively monocular maps of the central portions of the ipsilateral visual field in V1 and the contralateral visual field in V2. In addition, we document a complex interdigitation of V1 and V2 that entails a discontinuity in the mapping of visual space and fragmentation of V2 into isolated cortical territories. We suggest that both the monocularity of these cortical maps and the visuotopic discontinuity along the V1–V2 border derive from asymmetries in the crossed and uncrossed retinal pathways.

title: Orientation Selectivity and the Arrangement of Horizontal Connections in Tree Shrew Striate Cortex
abstract: Horizontal connections, formed primarily by the axon collaterals of pyramidal neurons in layer 2/3 of visual cortex, extend for millimeters parallel to the cortical surface and form patchy terminations. Previous studies have provided evidence that the patches formed by horizontal connections exhibit modular specificity, preferentially linking columns of neurons with similar response characteristics, such as preferred orientation. The issue of how these connections are distributed with respect to the topographic map of visual space, however, has not been resolved. Here we combine optical imaging of intrinsic signals with small extracellular injections of biocytin to assess quantitatively the specificity of horizontal connections with respect to both the map of orientation preference and the map of visual space in tree shrew V1. Our results indicate that horizontal connections outside a radius of 500 μm from the injection site exhibit not only modular specificity, but also specificity for axis of projection. Labeled axons extend for longer distances, and give off more terminal boutons, along an axis in the map of visual space that corresponds to the preferred orientation of the injection site. Inside of 500 μm, the pattern of connections is much less specific, with boutons found along every axis, contacting sites with a wide range of preferred orientations. The system of long-range horizontal connections can be summarized as preferentially linking neurons with co-oriented, co-axially aligned receptive fields. These observations suggest specific ways that horizontal circuits contribute to the response properties of layer 2/3 neurons and to mechanisms of visual perception.

title: Patterns of excitation and inhibition evoked by horizontal connections in visual cortex share a common relationship to orientation columns
abstract: Combined optical imaging and electrophysiological techniques were used to assess directly the functional nature of long-range excitatory and inhibitory synaptic interactions between orientation columns in area 17 of ferret visual cortex. A significant correlation was found between the layout of iso-orientation columns and the pattern of evoked synaptic inputs between cortical sites: the largest-amplitude inhibitory and excitatory synaptic responses were evoked in single neurons when stimulation and recording electrodes were located in orientation columns sharing the same angle preference. Both excitatory and inhibitory synaptic responses decreased in amplitude when stimulation and recording electrodes were located in orientation columns with orthogonal angle preferences. Changing the stimulus intensity altered the balance of evoked excitation and inhibition without changing the columnar specificity of inputs. These results directly demonstrate that horizontal connections modulate both excitatory and inhibitory synaptic interactions between iso-orientation columns.

title: Orientation selectivity, preference, and continuity in monkey striate cortex
abstract: Maps of orientation preference and selectivity, inferred from differential images of orientation (Blasdel, 1992), reveal linear organizations in patches, 0.5-1.0 mm across, where orientation selectivities are high, and where preferred orientations rotate linearly along one axis while remaining constant along the other. Most of these linear zones lie between the centers of adjacent ocular dominance columns, with their short iso-orientation slabs oriented perpendicular, in regions enjoying the greatest binocular overlap. These two-dimensional linear zones are segregated by one- and zero- dimensional discontinuities that are particularly abundant in the centers of ocular dominance columns, and that are also correlated with cytochrome oxidase-rich zones within them. Discontinuities smaller than 90 degrees extend in one dimension, as fractures, while discontinuities greater than 90 degrees are confined to points, in the form of singularities, that are generated when orientation preferences rotate continuously through +/- 180 degrees along circular paths. The continuous rotations through 180 degrees imply that direction preferences are not organized laterally in striate cortex. And they also ensure that preferences for all orientations converge at each singularity, with perpendicular orientations represented uniquely close together on opposite sides. The periodic interspersing of linear zones and singularities suggests that orientation preferences are organized by at least two competing schemes. They are optimized for linearity, along with selectivity and binocularity, in the linear zones, and they are optimized for density near singularities. Since upper-layer neurons are likely to have similarly sized dendritic fields in all regions (Lund and Yoshioka, 1991), those in the linear zones should receive precise information about narrowly constrained orientations, while those near singularities should receive coarse information about all orientations--very different inputs that suggest different perceptual functions.

title: Numerical Recipes in C: The Art of Sci-entific Computing
abstract: From the Publisher: 
This is the revised and greatly expanded Second Edition of the hugely popular Numerical Recipes: The Art of Scientific Computing. The product of a unique collaboration among four leading scientists in academic research and industry, Numerical Recipes is a complete text and reference book on scientific computing. In a self-contained manner it proceeds from mathematical and theoretical considerations to actual practical computer routines. With over 100 new routines (now well over 300 in all), plus upgraded versions of many of the original routines, this book is more than ever the most practical, comprehensive handbook of scientific computing available today. The book retains the informal, easy-to-read style that made the first edition so popular, with many new topics presented at the same accessible level. In addition, some sections of more advanced material have been introduced, set off in small type from the main body of the text. Numerical Recipes is an ideal textbook for scientists and engineers and an indispensable reference for anyone who works in scientific computing. Highlights of the new material include a new chapter on integral equations and inverse methods; multigrid methods for solving partial differential equations; improved random number routines; wavelet transforms; the statistical bootstrap method; a new chapter on "less-numerical" algorithms including compression coding and arbitrary precision arithmetic; band diagonal linear systems; linear algebra on sparse matrices; Cholesky and QR decomposition; calculation of numerical derivatives; Pade approximants, and rational Chebyshev approximation; new special functions; Monte Carlo integration in high-dimensional spaces; globally convergent methods for sets of nonlinear equations; an expanded chapter on fast Fourier methods; spectral analysis on unevenly sampled data; Savitzky-Golay smoothing filters; and two-dimensional Kolmogorov-Smirnoff tests. All this is in addition to material on such basic top

title: A Comparative Survey of Visual Cortex Organization in Mammals
abstract: A large region of posterior neocortex in a number of species of mammals has been found to be responsive to visual stimuli. In some of these mammals more detailed studies have subdivided the responsive region into separate systematic maps or representations of visual space, and there are several reasons for thinking that these maps are important as functionally distinct subdivisions of the visual system. For instance, it appears that each systematic map is associated with a cortical region having a unique architectonic character and a distinctive pattern of connections with other visual structures. In addition, there is evidence that neurons in separate visual representations differ in the way they respond to visual stimuli. Finally, lesions of different visual maps have produced distinctively different behavior changes.

title: Aberrant visual projections in the Siamese cat
abstract: 1. Guillery has recently shown that the Siamese cat has a grossly abnormal lateral geniculate body. His anatomical study suggested that certain fibres originating in the temporal retina of each eye cross in the chiasm instead of remaining uncrossed. They thus reach the wrong hemispheres, but in the geniculate they terminate in the regions that the missing fibres from the ipsilateral eye would normally have occupied. The result is that each hemisphere receives an input from parts of the ipsilateral field of vision, this input being entirely from the opposite eye. The purpose of the present work was to study the physiological consequences of this aberrant projection, in the lateral geniculate body and visual cortex.

title: A comparison of visual pathways in Boston and Midwestern Siamese cats
abstract: A genetic mutation in Siamese cats causes retinogeniculate fibers representing roughly the first 20 degrees of ipsilateral visual field in each eye to cross aberrantly in the optic chiasm and terminate in the wrong lateral geniculate nucleus (LGN). Previous investigations have shown that in the visual cortex this extra representation of ipsilateral visual field can be organized into one pattern in Boston Siamese cats, another in Midwestern. This finding was confirmed here.

title: Siamese cat: altered connections of visual cortex.
abstract: In siamese cats, each side of the brain receives a retinal input serving part of the ipsilateral visual field as well as the normal contralateral field representation. Both corticothalamic and cortico-cortical projections are systematically rearranged, but while one is retinotopically appropriate, the other fails to make a distinction between ipsilateral and contralateral fields. Different rules appear to govern the development of these two sets of connections.

title: Abnormal ipsilateral visual field representation in areas 17 and 18 of hypopigmented cats
abstract: We compared the central projections of retinal ganglion cells in temporal retina and the cortical representation of visual fields in areas 17 and 18 in cats with various hypopigmentation phenotypes (albino, heterozygous albino, Siamese, and heterozygous Siamese). In all cats studied, we found that the extent of abnormal ipsilateral visual field representation varied widely, and more of the ipsilateral visual field was represented in area 18 than in area 17. The greatest degree of ipsilateral visual field representation was found in albino cats, followed by Siamese, heterozygous albino and heterozygote Siamese cats, respectively. Additionally, in the different groups there was wide variation in the numbers of contralaterally projecting alpha and beta ganglion cells in temporal retina. In all cases, however, contralaterally projecting alpha cells were found to extend further into temporal retina than beta cells. We found that in each cat studied, the maximum extent of the abnormal ipsilateral visual field representation in areas 18 and 17 corresponded to the location of the 50% decussation line (i. e., the point where 50% of the ganglion cells in temporal retina project to the contralateral hemisphere) for alpha and beta cells, respectively, for that cat. Our results suggest that the extent of the abnormal visual field representations in visual cortex of hypopigmented cats reflects the extent of contralaterally projecting retinal ganglion cells in temporal retina.

title: The cortical visual areas of the sheep.
abstract: 1. A stereotaxic method for the sheep brain is described. 2. At its widest part the primary visual area (Visual I) of each hemisphere extends approximately 20 mm anteroposteriorly and, when unfolded, approximately 35 mm from side to side. It occupies both walls of the lateral sulcus, and extends medially to the medial wall of the hemisphere and to the depth of the ectolateral sulcus laterally. 3. The most lateral part of the primary visual area includes 10‐15 degrees of the ipsilateral field; the contralateral field is represented to 135 degrees from the mid line. 4. Visual II also includes a strip of ipsilateral representation on its medial edge and extends to the supra‐sylvian sulcus on the lateral surface of the brain. The furthest lateral representation recorded was 130 degrees lateral. 5. Most of both visual areas is concerned with the area centralis and the visual streak. The remainder of the retina has very little cortical representation. 6. Most cells in Visual I are simple with orientational and sometimes directional sensitivity. Some complex and hypercomplex cells have been seen in Visual I, and these predominate in Visual II. Receptive field sizes from 0‐25 to 10 degree were found. Within 15 degrees of the vertical meridian, binocular cells are common in both Visual I and II.

title: Functional organization in the visual cortex of the golden hamster
abstract: The visual cortex of the golden hamster was studied by means of multi‐unit and single unit recording, which revealed three separate retinotopic maps of the visual field in the posterior cortex. V1, corresponding to cytoarchitectonic area 17, has the contralateral temporal field represented medially, the central visual field (extending about 10 deg ipsilateral) represented laterally and the lower field anteriorly. The borders of the map, especially for the upper field, seem to be more restricted than the whole visual field available to the contralateral hemiretina: V1 probably does not represent the extreme periphery of the field. A large fraction of V1 has binocular input, for up to about 50 deg lateral to the vertical midline. There is a retinotopic reversal near the representation of the vertical midline where V1 meets V2 (corresponding to the more lateral “area 18a”). There is another retinotopic reversal at the extremity of the contralateral field representation, where V1 meets Vm (the medial visual area, corresponding to “area 18”) V2 and Vm each contain a reduced mirror image version of the map in V1. Almost all isolated single units in V1 have receptive fields that can be classified as radially symmetrical (60%) or asymmetrical (35%) Symmetrical fields have ON (13%) OFF (4%) ON‐OFF (30%) or “SILENT” (12%) central areas when plotted with flashing spots. There are minor but not striking differences between these groups in their field sizes, velocity preferences and so on. They almost invariably prefer moving to stationary stimuli but are not selective for orientation or direction of movement. Asymmetrical fields are of four types, three of which (type 1, 11% type 2, 17% and type 3, 2%) are orientation selective and resemble simple, complex and hypercomplex cells in the cat cortex. Some of these have direction as well as orientation preference. Axial movement detectors (5%) have a selectivity for one axis of motion, and thus prefer one orientation of edge, but respond equally well to movement of a spot. Vertical and horizontal orientation preferences, especially the latter, are much the most common. There is some evidence of clustering of cells according to receptive field type and, possibly, preferred orientation. Asymmetrical cells are, relatively, somewhat rarer in the deeper cortical layers. Within the binocular segment, fully 89%of cells are binocularly driven and the receptive fields are similar in the two eyes. Receptive fields tend to increase in size away from the area centralis representation and, in a complementary fashion, the magnification factor decreases from up to 0.1 mm/deg at the area centralis representation to about 0.02 mm/deg for the peripheral field.

title: The ipsilateral field representation in the striate cortex of the opossum
abstract: SummaryReference axes for the visuotopic study of the opossum's striate cortex were estimated from corresponding binocular response fields using multi-unit recording. These central binocular axes (CBA) were derived from experimental data based on the concept that corresponding receptive fields for each eye should be mostly in register under natural conditions. Vertical reference meridians, orthogonal to these axes, define a contralateral and an ipsilateral field for each eye with respect to the recording site. An ipsilateral field representation was observed for both eyes in the striate cortex at the transition zone with peristriate. Maximal values for the center and border of ipsilateral receptive fields were, respectively, 8 and 20 degrees for the contralateral eye and 6 and 14 degrees for the ipsilateral eye. An equivalent ipsilateral field representation was found in animals that had the anterior commissure cut prior to the recording session. This suggests that the ipsilateral field of both eyes may be represented in the striate cortex via the ipsilateral optic tract. Additionally, it was observed that the region of higher ganglion cell density in the retina shows a flattened distribution and that the CBA intersects the retina at the temporal aspect of this region.

title: The retinal projection to the thalamus in the cat: A quantitative investigation and a comparison with the retinotectal pathway
abstract: The projection of cat retinal ganglion cells to the thalamus was examined using the method of retrograde axonal transport of horseradish peroxidase (HRP). After the injection site was determined physiologically, HRP was applied by one of three methods: iontophoretic injection of minimal amounts, single pressure injections and multiple pressure injections.

title: Retinal decussation patterns in pigmented and albino ferrets
abstract: The decussation patterns of retinal ganglion cells in adult pigmented and albino ferrets were determined from the distribution of cells labelled after large unilateral injections of horseradish peroxidase into the visual pathway, involving the lateral geniculate nucleus and fibres of passage to the superior colliculus. About 6000 retinal ganglion cells project ipsilaterally in pigmented ferrets compared with only about 1500 in albino ferrets. In both strains, the vast majority of these cells (99 and 87% in pigmented and albino animals, respectively) are located in the temporal crescent, although we describe one albino ferret in which an aberrant uncrossed projection arises from nasal retina. In pigmented ferrets, there is a sharp nasotemporal division that runs through the area centralis; a small proportion of the ganglion cells in temporal crescent (less than 10%) does project contralaterally. In albinos, however, the majority of cells in temporal retina project contralaterally. There is no clear nasotemporal division in the albino retina; the density of uncrossed ganglion cells is reduced throughout temporal crescent and at no location exceeds the comparable density of the crossed projection. The peak density within the reduced uncrossed projection is also displaced away from the area centralis into temporal retina. Analysis of cell type on the basis of soma size indicates that whereas large horseradish peroxidase injections into the visual pathway of pigmented ferrets label all types of ganglion cell in the crossed projection, injections restricted to the superior colliculus label only those ganglion cells with large or small somata. The distribution of cell sizes in the crossed projection from temporal retina is biased towards small cells in the pigmented ferret but in albinos resembles that seen in the crossed projection from nasal retina. Thus the adult pigmented ferret has both a well developed nasotemporal division in which decussation lines are obvious in the crossed and uncrossed pathways and also, unlike rodents but like cats, a class of ganglion cell that does not project to the superior colliculus. The albino mutation both reduces the uncrossed projection throughout temporal retina, although the reduction is greatest close to the area centralis, and also commensurately increases the crossed projection from temporal retina.

title: Chiasmatic course of temporal retinal axons in the developing ferret
abstract: Recent studies on the distribution of optic axons in the mature visual pathways, as well as on the genesis of their ganglion cells of origin, suggest that the time of axonal arrival at the optic chiasm determines the side of the brain to which a temporal retinal axon will project. The present study has examined this issue directly in fetal ferrets, by determining the projection of the temporal retina at different developmental stages. Fetuses of known gestational age were fixed with paraformaldehyde and subsequently implanted with crystals of the carbocyanine dye, DiI, into eithe the temporal retina, or into one optic tract. The lipophilic diffusion of the dye within the plasma membrane of the axons revealed the course of temporal retinal fibers through the fetal chiasm, as well as the distribution of ganglion cells across the two retinae projecting to one optic tract.

title: First evidence of diversity in eutherian chiasmatic architecture: Tree shrews, like marsupials, have spatially segregated crossed and uncrossed chiasmatic pathways
abstract: In the optic chiasm of mammals, axons either cross the midline to the opposite side of the brain or remain uncrossed. In the eutherian species studied to date, uncrossed axons in the caudal nerve are found in all regions. In the chiasm, they are dispersed through the hemichiasm, with many axons approaching the midline and then turning back to enter the same side of the brain as the originating eye. In marsupials, by contrast, uncrossed axons never approach the midline; instead, they remain grouped in the lateral nerve and chiasm. The impression gained from these data is that there is a major difference in chiasmatic architecture between eutherian and marsupial mammals. Therefore, the mechanisms by which axons choose their route through the chiasm was also thought to differ between the two major groups of mammals. However, the present study shows that the chiasm of a highly visual eutherian mammal, the tree shrew, is similar to that found in marsupials, with uncrossed axons confined to lateral regions and not approaching the midline. However, unlike marsupials, in the tree shrew, optic fascicles in the chiasm are often separated by thick collagen bundles. It is probable that the chiasmatic structure described to date for eutherian mammals is not ubiquitous, as was previously thought, and theories explaining the mechanisms by which axons chose their route through the chiasm during development will have to be expanded.  1998.

title: Cortical connections of striate and extrastriate visual areas in tree shrews
abstract: 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.  1998.

title: Possible functions of the interhemispheric connexions between visual cortical areas in the cat.
abstract: The functions of interhemispheric axons linking the borders between cortical areas 17 and 18 on the two sides of the brain were investigated by two techniques. A well‐matched sample of neurones was recorded in the 17/18 border region before and after an extensive lesion was made in the corresponding part of the other hemisphere. The proportion of binocularly driven cells fell from 96% to 67%, confirming the results of Dreher & Cottee (1975). Orientation‐and direction‐selectivity, as well as the responsiveness of the population of neurones, seemed unaltered. The reduction in binocularity was much less convincing for cells in the body of area 17, even very close to the callosal‐recipient zone. Reversible cooling of the 17/18 border had no effect on the few cells recorded outside the callosal zone in the other hemisphere nor on eighteen of the thirty‐five cells recorded in the callosal zone. However, in ten cells the receptive field disappeared completely in one eye; in five cells there was a general reduction in responsiveness; two cells lost a portion of the receptive field, on the ipsilateral side, in both eyes. The receptive fields that were apparently transmitted via the corpus callosum lay around the vertical meridian of the visual field and were not restricted to the visual hemifield ipsilateral to the receiving hemisphere: their distribution overlapped that provided by the direct geniculo‐cortical input. The principal function of the callosal projection between the 17/18 borders may be to contribute to binocular convergence on cortical cells and perhaps to play a part in stereoscopic vision.

title: Visual receptive field properties of cells innervated through the corpus callosum in the cat
abstract: SummaryThe present experiment examined the receptive field (R.F.) properties of cortical cells which receive part of their input from the contralateral hemisphere via the corpus callosum. Two groups of cats were used for recording unit activity: a normal control group, and an experimental group consisting of cats which had their optic chiasmas split across the midline prior to the recording sessions. Acute recordings were carried out in the conventional manner using tungsten microelectrodes and N2O: O2 anaesthesia. The recording site was the 17–18 border. The stimulus consisted of a thin bar generated on an oscilloscope screen by a computer. The bar, whose orientation was varied automatically from 0 ° to 345 ° in 15 ° steps, was swept across the screen at constant speed orthogonal to the orientation. Various R.F. properties were studied using both quantitative and qualitative criteria. Thus, in the normal cat, simple, complex and hypercomplex type R.F.'s were found, whereas no callosally activated cell was of the simple type. The ocular dominance distribution found in the split chiasma cat was skewed towards the ipsilateral eye, although a fairly large number of cells could be driven with the two eyes. The R.F.'s of the callosally activated neurons were all situated close to the vertical meridian, which they sometimes straddled. Both in the normal and in the chiasma sectioned cats, the complex cells had larger R.F.'s than the other cell types. However, the R.F.'s determined through the ipsilateral eye was essentially of the same dimensions as those obtained through the indirect interhemispheric pathway, and this irrespective of cell type. Orientation specificity was similar for the two eyes in the split chiasma cats as it was for the normal cats although in the former the orientation tuning curve was narrower for the callosal pathway than for the more direct thalamo-cortical pathway. The results are interpreted within the context of the different functions ascribed to the corpus callosum in vision.

title: Tangential organization of callosal connectivity in the cat's visual cortex
abstract: 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.