A comparison of magnification functions in area 19 and the lateral suprasylvian visual area in the cat

A retinotopic map can be described by a magnification function that relates magnification factor to visual field eccentricity. Magnification factor for primary visual cortex (VI) in both the cat and the macaque monkey is directly proportional to retinal ganglion cell density. However, among those extrastriate areas for which a magnification function has been described, this is often not the case. Deviations from the pattern established in V1 are of considerable interest because they may provide insight into an extrastriate area's role in visual processing. The present study explored the magnification function for the lateral suprasylvian area (LS) in the cat. Because of its complex retinotopic organization, magnification was calculated indirectly using the known magnification function for area 19. Small tracer injections were made in area 17, and the extent of anterograde label in LS and in area 19 was measured. Using the ratio of cortical area labeled in LS to that in area 19, and the known magnification factor for area 19 at the corresponding retinotopic location, we were able to calculate magnification factor for LS. We found that the magnification function for LS differed substantially from that for area 19: central visual field was expanded, and peripheral field compressed in LS compared with area 19. Additionally, we found that the lower vertical meridian's representation was compressed relative to that of the horizontal meridian. We also examined receptive field size in areas 17, 19, and LS and found that, for all three areas, receptive field size was inversely proportional to magnification factor.     

Local precision of visuotopic organization in the middle temporal area (MT) of the macaque

Summary The representation of the visual field in the middle temporal area (MT) was examined by recording from single neurons in anesthetized, immobilized macaques. Measurements of receptive field size, variability of receptive field position (scatter) and magnification factor were obtained within the representation of the central 25°. Over at least short distances (less than 3 mm), the visual field representation in MT is surprisingly orderly. Receptive field size increases as a linear function of eccentricity and is about ten times larger than in V1 at all eccentricities. Scatter in receptive field position at any point in the visual field representation is equal to about one-third of the receptive field size at that location, the same relationship that has been found in V1. Magnification factor in MT is only about onefifth that reported in V1 within the central 5° but appears to decline somewhat less steeply than in V1 with increasing eccentricity. Because the smaller magnification factor in MT relative to V1 is complemented by larger receptive field size and scatter, the point-image size (the diameter of the region of cortex activated by a single point in the visual field) is roughly comparable in the two areas. On the basis of these results, as well as on our previous finding that 180° of axis of stimulus motion in MT are represented in about the same amount of tissue as 180° of stimulus orientation in V1, we suggest that a stimulus at one point in the visual field activates at least as many functional modules in MT as in V1.     

Saccade-related spread of activity across superior colliculus may arise from asymmetry of internal connections

The superior colliculus (SC) receives a retinotopic projection of the contralateral visual field in which the representation of the central field is expanded with respect to the peripheral field. The visual projection forms a nonlinear, approximately logarithmic, map on the SC. Models of the SC commonly assume that the function defining the strength of neuronal connections within this map (the kernel) depends only on the distance between two neurons, and is thus isotropic and homogeneous. However, if the connection strength is based on the distance between two stimuli in sensory space, the kernel will be asymmetric because of the nonlinear projection onto the brain map. We show, using a model of the SC, that one consequence of these asymmetric intrinsic connections is that activity initiated at one point spreads across the map. We compare this simulated spread with the spread observed experimentally around the time of saccadic eye movements with respect to direction of spread, differing effects of local and global inhibition, and the consequences of localized inactivation on the SC map. Early studies suggested that the SC spread was caused by feedback of eye displacement during a saccade, but subsequent studies were inconsistent with this feedback hypothesis. In our new model, the spread is autonomous, resulting from intrinsic connections within the SC, and thus does not depend on eye movement feedback. Other sensory maps in the brain (e.g., visual cortex) are also nonlinear and our analysis suggests that the consequences of asymmetric connections in those areas should be considered.     

Indirect, across-the-midline retinotectal projections and representation of ipsilateral visual field in superior colliculus of the cat

1. In agreement with previous work, we have found that the ipsilateral visual field is represented in an extensive rostral portion--from one-third to one-half--of the superior colliculus (SC) of the cat. This representation is binocular. The SC representation of the ipsilateral visual field can be mediated both directly, by crossed retinotectal connections originating from temporal hemiretina, and indirectly, by across-the-midline connections relaying visual information from one-half of the brain to contralateral SC. 2. In order to study the indirect, across-the-midline visual input to the SC, we have recorded responses of SC neurons to visual stimuli presented to either the ipsilateral or the contralateral eye of cats with a midsagittal splitting of the optic chiasm. Units driven by the ipsilateral eye, presumably through the direct retinotectal input and/or corticotectal connections from ipsilateral visual cortex, were found throughout the SC, except at its caudal pole, which normally receives fibers from the extreme periphery of the contralateral nasal hemiretina. Units driven by the contralateral eye, undoubtedly through an indirect across-the-midline connection, were found only in the anterior portion of the SC, in which is normally represented the ipsilateral visual field. Receptive fields in both ipsilateral and contralateral eye had properties typical of SC receptive fields in cats with intact optic pathways. 3. All units having a receptive field in the contralateral eye had also a receptive field in the ipsilateral eye; for each of these units, the receptive fields in both eyes invariably abutted the vertical meridian of the visual field. The receptive field in one eye had about the same elevation relative to the horizontal meridian and the same vertical extension as the receptive field in the other eye; the two receptive fields of each binocular unit matched each other at the vertical meridian and formed a combined receptive field straddling the vertical midline of the horopter...     

Effects of vision restoration training on early visual cortex in patients with cerebral blindness investigated with functional magnetic resonance imaging

Cerebral blindness is a loss of vision as a result of postchiasmatic damage to the visual pathways. Parts of the lost visual field can be restored through training. However, the neuronal mechanisms through which training effects occur are still unclear. We therefore assessed training-induced changes in brain function in eight patients with cerebral blindness. Visual fields were measured with perimetry and retinotopic maps were acquired with functional magnetic resonance imaging (fMRI) before and after vision restoration training. We assessed differences in hemodynamic responses between sessions that represented changes in amplitudes of neural responses and changes in receptive field locations and sizes. Perimetry results showed highly varied visual field recovery with shifts of the central visual field border ranging between 1 and 7°. fMRI results showed that, although retinotopic maps were mostly stable over sessions, there was a small shift of receptive field locations toward a higher eccentricity after training in addition to increases in receptive field sizes. In patients with bilateral brain activation, these effects were stronger in the affected than in the intact hemisphere. Changes in receptive field size and location could account for limited visual field recovery (± 1°), although it could not account for the large increases in visual field size that were observed in some patients. Furthermore, the retinotopic maps strongly matched perimetry measurements before training. These results are taken to indicate that local visual field enlargements are caused by receptive field changes in early visual cortex, whereas large-scale improvement cannot be explained by this mechanism.     

The representation of perceived angular size in human primary visual cortex

Two objects that project the same visual angle on the retina can appear to occupy very different proportions of the visual field if they are perceived to be at different distances. What happens to the retinotopic map in primary visual cortex (V1) during the perception of these size illusions? Here we show, using functional magnetic resonance imaging (fMRI), that the retinotopic representation of an object changes in accordance with its perceived angular size. A distant object that appears to occupy a larger portion of the visual field activates a larger area in V1 than an object of equal angular size that is perceived to be closer and smaller. These results demonstrate that the retinal size of an object and the depth information in a scene are combined early in the human visual system.     

Uptake of horseradish peroxidase by geniculo-cortical axons in the golden hamster: Analysis by computer reconstruction

Summary Micro-injections of horseradish peroxidase (HRP) were made into the visual cortex of the golden hamster. The projection lines of labelled neurons in the dorsal lateral geniculate nucleus (LGNd) were three-dimensionally reconstructed, using a computer graphics technique. The lines run rostrally and medially from their origins at the lateral surface of the nucleus. Using an anatomically determined retinotopic map of the LGNd, the positions of all labelled cells near the lateral surface were converted into equivalent visual field co-ordinates and displayed on a physiologically determined retinotopic map of the primary visual cortex. Comparison between the scatter of these equivalent retinotopic loci and an actual reconstruction of the injection site revealed that: 1. there was general agreement between the independent retinotopic maps of LGNd and visual cortex; 2. there was greater retinotopic scatter of labelled LGNd cells than could be accounted for by the area of tissue injury in the cortex; 3. the retinotopic scatter matched more closely the total visible halo of HRP staining in the grey matter; 4. HRP can be taken up from a cytoarchitectonic field into which it diffuses after injection into a neighbouring area; 5. HRP is probably not taken up by undamaged axons in the white matter. These results are compared with those obtained in other animals and other systems. No general rules emerge, but the possibility of uptake from wide areas of diffusion must be considered when interpreting results of HRP injection.Royal Society Locke Research Fellow, on leave from the Physiological Laboratory, Cambridge, England     

Distribution in areas 18 and 19 of neurons projecting to the pontine nuclei: a quantitative study in the cat with retrograde transport of HRP-WGA

Summary Following large injections of horseradish peroxidase — wheat germ agglutinin in the pontine nuclei, corticopontine neurons in areas 18 and 19 were quantitatively mapped and flat maps showing the distribution of retrogradely labeled cells were constructed. The areal borders were defined either cyto- and myeloarchitectonically or from standard retinotopic maps presented in frontal sections (Tusa et al. 1981). Maps of the retinotopic organization in areas 18 and 19 (Tusa et al. 1979) were transferred to the present flat maps. Thus, the number and distribution of pontine projecting cells could be correlated with the retinotopic organization. The cell density (number of labeled cells per mm² cortex) is in both areas highest in the cortex representing the lower and upper visual periphery and decreases towards the representation of the retinal central area. However, since in both areas 18 and 19 the visual field representation is twisted and portions of the visual field are magnified, the actual number of cells is higher in the cortex representing the central area and the lower medial visual field than in other parts. The cortex representing the lower hemifield contains approximately 2/3 (mean, N = 4) of the corticopontine cells in both areas. The average density of corticopontine cells increases from area 17 through 18 to 19, but the total number of cells within each of the areas is about the same (area 17 18000 cells, area 18 13400 cells, area 19 17200 cells; mean, N = 4; data on area 17 from Bjaalie and Brodal, 1983). In conclusion, areas 17, 18 and 19 contribute about equally in quantitative terms to the pontine nuclei. Furthermore, assuming that the corticopontine neurons transmit spatially relevant information, there is a moderate overrepresentation of central vision and the lower medial visual field in the pontine projection from areas 18 and 19. This visual field representation is remarkably similar to that found in the corticopontine projection from area 17 (Bjaalie and Brodal 1983).     

The effects of prenatal and neonatal monocular enucleation on visual topography in the uncrossed retinal pathway to the rat superior colliculus

Summary The visual representation in the uncrossed retinal projection to the superior colliculus (SC) was examined electrophysiologically by recording multiunit responses in paralysed, anaesthetised adult rats (both pigmented and albino), which had been monocularly enucleated either prenatally or soon after birth. This manipulation partially stabilises an exuberant neonatal projection from the remaining eye to the ipsilateral SC. Neuronal responses were also stronger and the multi-unit receptive fields larger than in intact animals. Many of the visual fields recorded on penetrations in caudal SC were located in the peripheral ipsilateral visual hemifield, corresponding to nasal retina. Such receptive fields are not seen in normal animals and were not found in animals enucleated on day 3 or later. The topographic representation of the dorso-ventral retinal axis, lateral to medial in the SC, was normal in all experimental animals. The representation of the naso-temporal retinal axis was abnormal and more variable. In all operated animals as the recording electrode was moved caudally away from the rostral pole of the SC, the corresponding receptive fields moved gradually from up to 40° in the ipsilateral visual hemifield to about 40° into the contralateral hemifield (a location corresponding to the peripheral edge of the temporal retina). This is the mapping polarity found in the normal uncrossed retinal projection. In the enucleated animals, the map was expanded and frequently displayed a clustering of fields arising from far temporal retina. In animals enucleated prenatally or on the day of birth, visual responses could be recorded in more caudal SC. The corresponding receptive fields now moved nasally on the retina, generating reversals in the map. The most caudal penetrations in these early enucleates frequently gave receptive fields located in retina nasal to the optic disc, up to 90 degrees into the ipsilateral visual hemifield. These results demonstrate that a temporal relationship exists between the order and mapping polarity of the visual field in SC and the time of enucleation. Prenatal enucleation produces reversals of the mapping polarity in caudal SC while neonatal enucleation produces an expanded map but one with a mapping polarity appropriate for an uncrossed projection     

Spatial relation of the acetylcholinesterase-rich domain to the visual topography in the feline superior colliculus

Summary The superior colliculus (SC) of the cat shows a prominent compartmentalized organization at the level of its intermediate layers. The mosaic of these compartments is apparent in the pattern of acetylcholinesterase (AChE) staining. Patches of high AChE-activity are sharply set off from surrounding areas in the caudal SC while they are less distinct anteriorly. The rostral part lacks such obvious compartments. Thus, a structural reorganization apparently cuts across the topographical representations spread out in the SC. In order to test if this compartmental gradient relates to the topographic maps of the colliculus, retinotopic landmarks were visualized in the superficial layers by labeling the retinotectal pathway. In the SC ipsilateral to the eye injected with horseradish peroxidase (HRP) a paucity of labeling indicated the zone representing the ipsilateral visual half-field. Serial reconstructions of collicular sections, cut longitudinally or tangentially, revealed that the non-compartmentalized part of the intermediate layers corresponds to the representation of the ipsilateral visual half-field in the layers above, while an intricate mosaic array of compartments prevail in tectal zones related to the representation of the contralateral visual half-field.     

NMDA antagonists in the superior colliculus prevent developmental plasticity but not visual transmission or map compression

Partial ablation of the superior colliculus (SC) at birth in hamsters compresses the retinocollicular map, increasing the amount of visual field represented at each SC location. Receptive field sizes of single SC neurons are maintained, however, preserving receptive field properties in the prelesion condition. The mechanism that allows single SC neurons to restrict the number of convergent retinal inputs and thus compensate for induced brain damage is unknown. In this study, we examined the role of N-methyl-D-aspartate (NMDA) receptors in controlling retinocollicular convergence. We found that chronic 2-amino-5-phosphonovaleric acid (APV) blockade of NMDA receptors from birth in normal hamsters resulted in enlarged single-unit receptive fields in SC neurons from normal maps and further enlargement in lesioned animals with compressed maps. The effect was linearly related to lesion size. These results suggest that NMDA receptors are necessary to control afferent/target convergence in the normal SC and to compensate for excess retinal afferents in lesioned animals. Despite the alteration in receptive field size in the APV-treated animals, a complete visual map was present in both normal and lesioned hamsters. Visual responsiveness in the treated SC was normal; thus the loss of compensatory plasticity was not due to reduced visual responsiveness. Our results argue that NMDA receptors are essential for map refinement, construction of receptive fields, and compensation for damage but not overall map compression. The results are consistent with a role for the NMDA receptor as a coincidence detector with a threshold, providing visual neurons with the ability to calculate the amount of visual space represented by competing retinal inputs through the absolute amount of coincidence in their firing patterns. This mechanism of population matching is likely to be of general importance during nervous system development.     

Differences of visual field representation in the medial and lateral banks of the suprasylvian cortex (PMLS/PLLS) of the cat

Summary We have studied the orderliness of representation of visual space in the medial and lateral banks of the middle suprasylvian sulcus. Penetrations were made either parallel to the sulcus, in one bank or the other, or vertical, thus crossing the sulcus between the postero-medial (PMLS) and posterolateral (PLLS) divisions of this area. In some cases we found clear evidence for topographical order in the representation of the visual field with a tendency (greater in PMLS than in PLLS) for the receptive fields of cells recorded deeper in the walls of the sulcus to lie closer to the area centralis, but along many penetrations the receptive fields were so large and so scattered that no retinotopic arrangement could be discerned. In PMLS the receptive fields of the majority of units we studied were centred below and close to the horizontal meridian, whereas in PLLS they were distributed over both the upper and lower visual fields with an over-representation of the upper field. Receptive fields were significantly larger in PLLS (mean field area = 442.2 deg²) than in PMLS (mean area = 154.4 deg²); there was also less clear correlation between receptive field size and eccentricity in PLLS (correlation coefficient = +0.25) than in PMLS (corr. coeff. = +0.72). Analysis of the distance between the receptive field centres of consecutively recorded units demonstrated that the mean scatter in both PMLS and PLLS amounts to about half the average receptive field diameter. In summary the topographical representation of visual space is less orderly in PLLS, and may involve a wider area of the visual field. These findings may relate to the segregated visual cortical and extrageniculate thalamic connections that the medial and lateral banks of the LS receive.     

Uniform spatial spread of population activity in primate parafoveal V1

What are the shape and size of the region in primate V1 that processes information from a single point in visual space? This region, a fundamental property termed cortical point image (CPI) (McIlwain 1986), represents the minimal population of V1 neurons that can be activated by a visual stimulus and therefore has important implications for population coding in the cortex. Previous indirect attempts to measure the CPI in macaque V1 using sparse microelectrode recordings resulted in conflicting findings. Whereas some early studies suggested that CPI size is constant throughout V1 (e.g., Hubel and Wiesel 1974), others have reported large changes in CPI size in parafoveal V1 (e.g., Van Essen et al. 1984). To resolve this controversy, we used voltage-sensitive dye imaging in V1 of fixating monkeys to directly measure the subthreshold CPI and several related properties across a range of parafoveal eccentricities. We found that despite large changes in other properties of the retinotopic map, the subthreshold CPI is approximately constant and extends over ～6 × 8 mm(2). This large and invariant CPI ensures a uniform representation of each point in visual space, with a complete representation of all visual features in V1, as originally proposed by Hubel and Wiesel (1974). In addition, we found several novel and unexpected asymmetries and anisotropies in the shapes of the CPI and the population receptive field. These results expand our understanding of the representation of visual space in V1 and are likely to be relevant for the representations of stimuli in other sensory cortical areas.     

Responses and properties of receptive fields of neurons in the visual projection zone of the pigeon hyperstriatum

Unit responses in the hyperstriatal region of the pigeon forebrain to the action of various visual stimuli were investigated. Particular attention was paid to the discovery of retinotopic projection in the Wulst region. It was shown that as the electrode was advanced in the caudal direction in the zone of visual projection of the hyperstriatum the receptive fields of the neurons recorded shifted in the opposite direction in the visual field. The receptive fields of neurons of the ventral and dorsal hyperstriatum lie higher in the visual field and are larger in diameter than those of neurons of the accessory hyperstriatum. Unit responses in the visual projection zone of the Wulst depend on the intensity of illumination, size, and speed and direction of movement of the test objects across the receptive field. The functional role of the retino-thalamo-telencephalic system in visual interpretation in birds is discussed and it is suggested that the Wulst region is comparable with the striatal and also with the frontal regions of the mammalian cortex.Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 230–236, May–June, 1976.     

Visual responses of the human superior colliculus: a high-resolution functional magnetic resonance imaging study

The superior colliculus (SC) is a multimodal laminar structure located on the roof of the brain stem. The SC is a key structure in a distributed network of areas that mediate saccadic eye movements and shifts of attention across the visual field and has been extensively studied in nonhuman primates. In humans, it has proven difficult to study the SC with functional MRI (fMRI) because of its small size, deep location, and proximity to pulsating vascular structures. Here, we performed a series of high-resolution fMRI studies at 3 T to investigate basic visual response properties of the SC. The retinotopic organization of the SC was determined using the traveling wave method with flickering checkerboard stimuli presented at different polar angles and eccentricities. SC activations were confined to stimulation of the contralateral hemifield. Although a detailed retinotopic map was not observed, across subjects, the upper and lower visual fields were represented medially and laterally, respectively. Responses were dominantly evoked by stimuli presented along the horizontal meridian of the visual field. We also measured the sensitivity of the SC to luminance contrast, which has not been previously reported in primates. SC responses were nearly saturated by low contrast stimuli and showed only small response modulation with higher contrast stimuli, indicating high sensitivity to stimulus contrast. Responsiveness to stimulus motion in the SC was shown by robust activations evoked by moving versus static dot stimuli that could not be attributed to eye movements. The responses to contrast and motion stimuli were compared with those in the human lateral geniculate nucleus. Our results provide first insights into basic visual responses of the human SC and show the feasibility of studying subcortical structures using high-resolution fMRI.     

Unusually large receptive fields in cats with restricted visual experience

Summary The receptive fields of striate cortex neurons were analyzed in cats which had restricted or no visual experience. Two groups of animals were investigated: 1. cats which were deprived from contour vision over variable periods of time up to 1 year and 2. kittens whose visual experience was restricted to vertically oriented gratings of constant spatial frequency which moved unidirectionally at a fixed distance in front of the restrained animals. In both preparations exceedingly large receptive fields (up to 20° in diameter) were encountered, especially in cells located in supragranular layers. These large receptive fields never extended over more than 2° into the ipsilateral hemifield. Their sensitivity profile was frequently asymmetric and contained discontinuities. Many of these large receptive fields consisted of several excitatory subregions which were separated from each other by as much as 15°. Often but not always the most sensitive area was located where the retinotopic map predicted the receptive field center. The orientation and direction selectivity and also the angular separation of such multiple excitatory bands often matched precisely the orientation, direction and spatial frequency of the experienced moving grating. In other fields with multiple excitatory subregions such a correspondence could not be established; the various subregions could even have different orientation and direction selectivities. From these unconventional receptive fields it is concluded that the function of cat striate cortex is not confined to a point by point analysis of the visual field in retinotopically organized and functionally isolated columns.     

Effects of cooling somatosensory cortex on response properties of tactile cells in the superior colliculus

The corticotectal influences of somatosensory cortex were investigated by using reversible deactivation of cortex by cooling. More than half of the somatosensory superior colliculus (SC) cells studied exhibited a response depression (often not apparent qualitatively) or an elimination of responses to somatosensory stimuli during the period in which cortex was rendered inactive. Responses were restored to their initial levels by cortical rewarming. Hyperresponsiveness was never observed as a consequence of cortical cooling. Susceptibility to cooling-induced depression was not invariably linked to a specific cell type, location in the SC, or receptive-field size. Yet cells that had small receptive fields and were activated by hair displacement had the highest probability of being affected by this procedure. In some cells a contraction of the receptive field was induced by cortical cooling. This observation is consistent with previous experiments that showed that SC somatosensory receptive fields are constructed by the convergence of ascending and descending inputs and indicates that the responsiveness of specific receptive-field regions may depend on the functional integrity of cortex. Two cortical regions were found to produce cooling-induced effects in somatosensory SC cells: 1) SIV (and para-SIV), located in the anterior ectosylvian sulcus, and 2) the cortex within the rostral suprasylvian sulcus. These results indicate that somatosensory cortex, like visual cortex, plays a critical role in modulating the responses of SC cells. Apparently, the ability of both somatosensory and visual SC cells to code the presence of peripheral stimuli depends largely on the functional influences of their respective cortices. However, in contrast to previous observations on visual corticotectal influences, no specific receptive-field properties could be shown to be impressed on SC cells by somatosensory cortex.     

Retinotopic organization of extra-retinal saccade-related input to the visual cortex in the cat

Summary Single unit activity of 842 cells has been recorded in cat visual cortex and analyzed with respect to vestibular induced, and spontaneous saccadic eye movements performed in the dark. This study has been done in awake, chronically implanted cats, subsequently placed in acute conditions to achieve the precise retinotopic mapping of the cortical areas previously investigated.In areas 17 and 18, respectively, 27% and 24% of the cells tested were influenced by horizontal saccadic eye movements in the dark (E. M. cells). In the Clare-Bishop area, the proportion of E. M. cells was 12%, while only 2% of such cells were found in areas 19 and 21.The distribution of E.M. cells in areas 17 and 18 with respect to retinotopy showed that E.M. cells were more numerous in the cortical zones devoted to the representation of the area centralis (38% in area 17, 27% in area 18) than in the zones subserving the periphery of the visual field (17% and 12%, respectively).Two of the characteristics of E. M. cell activations appear dependant on the retinotopic organization. First, a larger number of E.M. cells presenting an asymmetry in their responses to horizontal saccadic eye movements in opposite directions (directional E.M. cells) were encountered in the cortical representation of the peripheral visual field. 53% of E. M. cells recorded in area 17 and 71% in area 18 were directional in the cortex corresponding to the peripheral visual field. This percentage was of 23% and 25% respectively in the cortex devoted to area centralis. Second, E.M. cells were found to have a latency from the onset of the saccade systematically larger than 100 ms (i.e, they discharged at, or after the end of the eye movement) if they were located in the cortical representation of the area centralis, while E.M. cells related to the peripheral visual field displayed a wider range of latencies (0–240 ms).Results obtained in Clare Bishop area, although limited to the representation of the peripheral visual field, were quantitatively and qualitatively similar to those observed in the homologous retinotopic zones of areas 17 and 18.It is concluded that an extra-retinal input related to oculomotor activity is sent to the cat visual cortex and is organized, at least in areas 17 and 18, with respect to the retinotopic representation of the visual field. These data support the hypothesis of a functional duality between central and peripheral vision and are discussed in the context of visual-oculomotor integration.Supported by INSERM (C.R.L. 79-53336)     

Interhemispheric influences on area 19 of the cat

Summary Anatomical studies have shown an extensive network of homotopic and heterotopic interhemispheric connections in area 19 of the cat visual cortex (Segraves and Rosenquist 1982a; 1982b). We have investigated their functional organization by recording visual responses in area 19 of cats following a midsagittal section of the optic chiasm. This operation interrupts all crossed optic fibers coming both from the nasal and the temporal retinae; as a result, each hemisphere receives optic fibers only from the lateral hemiretina of the ipsilateral eye which conveys information from the contralateral visual field. Visual information transmitted to the same hemisphere from the contralateral retina and the ipsilateral visual field must be attributed to an indirect, interhemispheric pathway. We found that a rather high proportion of neurons (31.8%) in area 19 of seven split-chiasm cats responded to visual stimuli presented to the contralateral eye. 1 — All neurons receiving this interhemispheric activation were also driven by the ipsilateral eye via an intrahemispheric pathway. 2 — The property of binocularity was significantly related to the visuotopic map in that both receptive fields of each binocular neuron adjoined or were in the immediate vicinity of the vertical meridian. 3 — Due to the small size of receptive fields in area 19, the contribution of the interhemispheric pathway to the representation of the visual field is rather limited and it is certainly less extensive than that predicted by anatomical studies. The representation of the ipsilateral visual field in area 19 of intact cats, as assessed electrophy-siologically, was comparable to that found in split-chiasm cats. Recordings in areas 17–18 of split-chiasm cats showed that the visual field represented through the corpus callosum in these visual areas is certainly not less and probably more, extensive than that found in area 19. The results support the conclusion that the relation to the vertical meridian and the receptive field size can explain the organization of the interhemispheric connections in the visual areas studied so far.     

Simultaneous constraints on pre- and post-synaptic cells couple cortical feature maps in a 2D geometric model of orientation preference.

The most prominent feature of mammalian striate cortex (V1) is the spatial organization of response preferences for the position and orientation of elementary visual stimuli. Models for the formation of cortical maps of orientation and 'retinotopic' position typically rely on a combination of Hebbian or correlation-based synaptic plasticity, and constraints on the distribution of synaptic weights. We consider a simplified model of orientation and retinotopic specificity based on the geometry of the feed-forward synaptic weight distribution from an 'unoriented' layer of cells to a first weakly oriented layer. We model the feed-forward weight distribution as a system of planar Gaussian receptive fields each elongated in the direction matching the preferred orientation of the postsynaptic cell. Under the constraint of presynaptic weight normalization (each cell in the oriented layer receives the same net synaptic weight) and a uniform retinotopic map (displacement of centres of mass of receptive fields in the unoriented layer is strictly proportional to the displacement of the corresponding cells in the oriented layer), we find that imposing a pattern of orientation preference forces the system to violate postsynaptic weight normalization (each cell in the unoriented layer no longer sends forth the same net synaptic weight). We study this deviation from uniformity of the postsynaptic weight, and find that the deviation has a distinct form in the vicinity of the 'pinwheel' singularities of the orientation map. We show that uniform synaptic coverage of the unoriented layer can be restored by introducing a distortion in the retinotopic locations of the receptive fields. We calculate, to first order in the relative elongation of the receptive fields, the retinotopic distortion vector field. Both the pattern of postsynaptic weight non-uniformity and the corrective retinotopic distortion vector field fail to possess the reflection symmetry commonly assumed to relate orientation singularities with topological index +/- pi. Hence, we show that 'right-handed' and 'left-handed' orientation singularities are funda-mentally distinct anatomical structures when full 2D synaptic architecture is taken into account. Finally, we predict specific patterns of retinotopic distortion that should obtain in the vicinity of +/- pi-fold orientation singularities, if uniform pre- and post-synaptic weight constraints are strongly enforced.