Lateral Suprasylvian Visual Area Research Articles

Cortical and thalamic connectivity of occipital visual cortical areas 17, 18, 19, and 21 of the domestic ferret (Mustela putorius furo).

The present study describes the ipsilateral and contralateral corticocortical and corticothalamic connectivity of the occipital visual areas 17, 18, 19, and 21 in the ferret using standard anatomical tract-tracing methods. In line with previous studies of mammalian visual cortex connectivity, substantially more anterograde and retrograde label was present in the hemisphere ipsilateral to the injection site compared to the contralateral hemisphere. Ipsilateral reciprocal connectivity was the strongest within the occipital visual areas, while weaker connectivity strength was observed in the temporal, suprasylvian, and parietal visual areas. Callosal connectivity tended to be strongest in the homotopic cortical areas, and revealed a similar areal distribution to that observed in the ipsilateral hemisphere, although often less widespread across cortical areas. Ipsilateral reciprocal connectivity was observed throughout the visual nuclei of the dorsal thalamus, with no contralateral connections to the visual thalamus being observed. The current study, along with previous studies of connectivity in the cat, identified the posteromedial lateral suprasylvian visual area (PMLS) as a distinct network hub external to the occipital visual areas in carnivores, implicating PMLS as a potential gateway to the parietal cortex for dorsal stream processing. These data will also contribute to a macro connectome database of the ferret brain, providing essential data for connectomics analyses and cross-species analyses of connectomes and brain connectivity matrices, as well as providing data relevant to additional studies of cortical connectivity across mammals and the evolution of cortical connectivity variation.

Congenital Deafness Reduces, But Does Not Eliminate Auditory Responsiveness in Cat Extrastriate Visual Cortex

Congenital deafness not only affects the development of the auditory cortex, but also the interrelation between the visual and auditory system. For example, congenital deafness leads to visual modulation of the deaf auditory cortex in the form of cross-modal plasticity. Here we asked, whether congenital deafness additionally affects auditory modulation in the visual cortex. We demonstrate that auditory activity, which is normally present in the lateral suprasylvian visual areas in normal hearing cats, can also be elicited by electrical activation of the auditory system with cochlear implants. We then show that in adult congenitally deaf cats auditory activity in this region was reduced when tested with cochlear implant stimulation. However, the change in this area was small and auditory activity was not completely abolished despite years of congenital deafness. The results document that congenital deafness leads not only to changes in the auditory cortex but also affects auditory modulation of visual areas. However, the results further show a persistence of fundamental cortical sensory functional organization despite congenital deafness.

Amplified somatosensory and visual cortical projections to a core auditory area, the anterior auditory field, following early- and late-onset deafness.

Cross-modal reorganization following the loss of input from a sensory modality can recruit sensory-deprived cortical areas to process information from the remaining senses. Specifically, in early-deaf cats, the anterior auditory field (AAF) is unresponsive to auditory stimuli but can be activated by somatosensory and visual stimuli. Similarly, AAF neurons respond to tactile input in adult-deafened animals. To examine anatomical changes that may underlie this functional adaptation following early or late deafness, afferent projections to AAF were examined in hearing cats, and cats with early- or adult-onset deafness. Unilateral deposits of biotinylated dextran amine were made in AAF to retrogradely label cortical and thalamic afferents to AAF. In early-deaf cats, ipsilateral neuronal labeling in visual and somatosensory cortices increased by 329% and 101%, respectively. The largest increases arose from the anterior ectosylvian visual area and the anterolateral lateral suprasylvian visual area, as well as somatosensory areas S2 and S4. Consequently, labeling in auditory areas was reduced by 36%. The age of deafness onset appeared to influence afferent connectivity, with less marked differences observed in late-deaf cats. Profound changes to visual and somatosensory afferent connectivity following deafness may reflect corticocortical rewiring affording acoustically deprived AAF with cross-modal functionality.

Evidence regarding the integrity of the posterior medial lateral suprasylvian visual area in the cat

Among the areas of lateral suprasylvian visual cortex in cats defined by Palmer et al. (J Comp Neurol [1978] 177:237-256), PMLS (posterior lateral suprasylvian area) has been the most studied. Although PMLS has strong and well-documented connections with area 17, it is unclear whether these connections extend to its upper visual field representation. We asked what cortical areas send input to the upper field representation in PMLS by making tracer injections in areas 17, 19, and posterior suprasylvian cortex. Tracer injections made in area 17's upper field representation in 15 cats failed to label the corresponding region in PMLS. Instead, they showed that area 17 is strongly connected with the posterior bank of the posterior suprasylvian sulcus (pSS), a region attributed by Palmer et al. to area 21a. Injections in area 19 had the same outcome. We consider this posterior upper field representation plus the lower field representation in PMLS to belong to a single area, LS (lateral suprasylvian visual area). Our data suggest that the upper field representation in PMLS belongs to a different area, most likely AMLS (anterior medial lateral suprasylvian area).

Location, architecture, and retinotopy of the anteromedial lateral suprasylvian visual area (AMLS) of the ferret (Mustela putorius)

The present paper describes the results of architectural and electrophysiological mapping observations of the medial bank of the suprasylvian sulcus of the ferret immediately caudal to somatosensory regions. The aim was to determine if the ferret possessed a homologous cortical area to the anteromedial lateral suprasylvian visual area (AMLS) of the domestic cat. We studied the architectural features and visuotopic organization of a region that we now consider to be a homologue to the cat AMLS. This area showed a distinct architecture and retinotopic organization. The retinotopic map was complex in nature with a bias towards representation of the lower visual field. These features indicate that the region described here as AMLS in the ferret is indeed a direct homologue of the previously described cat AMLS and forms part of a hierarchy of cortical areas processing motion in the ferret visual cortex. With the results of the present study and those of earlier studies a total of twelve cortical visual areas have been determined presently for the ferret, all of which appear to have direct homologues with visual cortical areas in the cat (which has a total of eighteen areas).

Responses in extrastriate cortex to optic flow during simulated turns.

An observer locomoting along a straight path sees a pattern of optic flow in which images move approximately radially outward from his heading point. If the observer turns, the optic flow field changes markedly because each object's image now has a component of horizontal motion added to its "optic flow" motion. We tested the responses of 326 cells in the cat's extrastriate area LS (lateral suprasylvian visual area) to movies simulating the optic flow seen during locomotion in a straight line, and during various simulated turns to the left and right. About 60% of 326 cells tested responded to optic flow simulating turns. Of most interest was a subset of cells, 15% of the total, that had "turn-selective" responses. They responded significantly better to turns in a particular direction (usually to the contralateral side) than to turns in the opposite direction or to optic flow simulating straight-ahead locomotion. For each cell, we generated a display of fronto-parallel motion with a direction and speed that matched the image motion in the preferred turn movie, as seen at the receptive-field center. Most turn-selective cells responded significantly better to their preferred turn movie than to this fronto-parallel stimulus. We examined the role of cells' selectivity for stimulus direction, speed, and acceleration in determining cell preference for particular turns. Direction preference played some role for most cells, but about a third of the cells preferred turn movies that did not reflect their direction selectivity. Other factors, including the presence of opposing motion, only rarely appeared to determine cell preferences for particular turn movies.

Lesions of extrastriate cortex and consequences for visual guidance during locomotion.

One of the functions of the putative motion pathway in visual cortex may be visual guidance during locomotion. Using cats, we have investigated the role of the first area in this pathway, the lateral suprasylvian visual area (LS, often considered analogous to the primate area MT). Visual function during locomotion was tested by looking at the accuracy of foot placement as cats walked down a cluttered alley. Cats were tested before and after LS was lesioned bilaterally using ibotenic acid. In all four cats tested, partial lesions of LS caused a significant deterioration in the accuracy of foot placement, though cats showed no deficit in acuity or in pattern discrimination. To determine whether the observed deficit was specific to the motion pathway, in two additional cats lesions were made in the central field representation in area 19, which lies adjacent to LS but belongs to the putative object recognition pathway. Surprisingly, both cats showed significant improvements in their accuracy of foot placement during locomotion. The reason for this improvement was unclear.

Effects of Superior Colliculus Inhibition on Three-Dimensional Visual Motion Processing in the Lateral Suprasylvian Visual Area of the Cat

Purpose: To determine whether visual inputs from the tectothalamocortical pathway influence three-dimensional motion processing within the lateral suprasylvian (LS) area of the cat. Methods: Tungsten microelectrodes were used for recording visual-evoked potentials (VEPs) from the LS area of 4 cats. Random dot stereograms were used as visual stimuli. Three-dimensional, motion-triggered VEPs were recorded from the LS area. Each motion sequence consisted of an abrupt onset of motion disparity with a 2° amplitude followed by an abrupt offset and a stationary phase of 900 ms. The velocity of the motion disparity was varied in eight steps from 10° to 400° per second. The onset of motion disparity was used as the trigger for recording the VEPs. Single or multiple injections (two to three) of muscimol were made mainly into the rostral superior colliculus (SC). The amplitudes of the VEPs before and after the muscimol injection were compared. Results: A large negative wave ( N1) with an implicit time of 92.7 ± 13.5 ms (mean ± SD, n = 98) was recorded consistently. The amplitude of N1 was significantly larger on stereovision of motion disparity than on either binocular vision of two-dimensional lateral motion or monocular vision, indicating that N1 contains neurons sensitive to motion disparity. The amplitude of N1 was not altered by muscimol injection into the SC at velocities ≤50°/s. On the other hand, the amplitude of N1 was reduced to 66–71% of that observed before muscimol injection at velocities ≥75°/s. Conclusions: These findings suggest that the LS area processes three-dimensional motion inputs via two parallel pathways, the geniculostriate pathway and the tectothalamocortical pathway, at velocities of motion disparity ≥75°/s, while the three-dimensional motion inputs project to the LS area only via the geniculostriate pathway at velocities of motion disparity ≤ 50°/s.

Two visual areas located in the middle suprasylvian gyrus (cytoarchitectonic field 7) of the cat's cortex

Neuronal properties and topographic organization of the middle suprasylvian gyrus (cortical cytoarchitectonic field 7) were studied in three behaving cats with painlessly fixed heads. Two main neuronal types were found within this field. Type 1 neurons occupied the lateral part of the field and bordered representation of directionally selective neurons of the lateral suprasylvian visual area by vertical retinal meridian. Type 1 neurons had elongated and radially oriented receptive fields located in the lower part of contralateral visual field. Type 1 neurons preferred stimuli moving out or to the centre of gaze at a low or moderate speed, and many of them were depth selective. The responses were enhanced by attention, oriented to the presented stimulus. Medial part of the field 7 along the border with the area V3 was occupied by neurons with not elongated receptive fields (type 2). These neurons preferred moderate and high speeds of motion, and gratings of proper spatial frequency and orientation were effective stimuli for them. Border between representations of type 2 and type 1 neurons coincided with projection of horizontal retinal meridian. At the rostral and caudal borders of the field 7 abrupt changes of neuronal properties took place. Neurons which abutted field 7 anteriorly and posteriorly resembled hypercomplex cells and their small receptive fields were located in the central part of the visual field. Topographical considerations and receptive field properties allowed us to conclude that the medial part of the field 7 (included type 2 neurons) is functionally equivalent to the area V4 in the cortex of primates, while the lateral part (type 1 neurons) may correspond to the area V4T.

Thalamic control of cat area-18 supragranular layers: simple cells, complex cells, and cells projecting to the lateral suprasylvian visual area.

The goal of this study was to determine the effects of inactivating layer A or the C layers of the cat lateral geniculate nucleus on the supragranular layers of area 18, including cells antidromically activated from the lateral suprasylvian visual area (LS). Isolated cells were visually driven via the contralateral eye while the retinotopically corresponding regions of layer A or, in some cases, the C layers were reversibly inactivated with injections of cobaltous chloride. Simple cells were frequently encountered and were on average more dependent upon layer A than were complex cells, a result qualitatively similar to that found previously in area 17 (Malpeli, 1983; Malpeli et al., 1986). However, the influence of the C layers on area 18 was much more apparent than for area 17. In area 18, as in area 17, the dependence of simple cells on particular geniculate layers appears to follow the terminal patterns of the major direct geniculate inputs. Those simple cells most dependent on layer A were located in lower layer 3. Simple cells in upper layer 3, like complex cells, showed little dependence on layer A, but were strongly dependent upon the C layers. All cells antidromically activated from LS were simple cells with rapidly conducting axons. They had, on average, the same moderately strong dependence on layer A as the patches of LS receiving area 18 input (Lee et al., 1997), supporting the conclusion that the influence of layer A in these patches is largely transmitted via association inputs from area 18. These results demonstrate that simple cells play a major role in association pathways.

Thalamic control of cat lateral suprasylvian visual area: relation to patchy association projections from area 18.

In this study, we examined functional contributions of major subdivisions of the lateral geniculate nucleus to the cat's lateral suprasylvian visual area (LS) in relation to the patchy horizontal distributions of association inputs. Multiple-unit activity driven via the contralateral eye was assessed during reversible blockade of the retinotopically corresponding part of layer A, the C layers as a group, or the medial interlaminar nucleus (MIN). Inactivating each of these targets reduced activity at some cortical sites, with inactivation of layer A having, on average, the largest effect. Activity was rarely abolished by inactivation of a single target, indicating that most LS sites receive multiple inputs. Dependence on layer A was strongly correlated with the horizontal distribution of association inputs from area 18. Closely spaced injections of anatomical tracers into extensive regions of area 18 resulted in patches of terminal label in lateral suprasylvian cortex. Activity inside the patches was relatively dependent on layer A, whereas that outside the patches was not. Dependence on the MIN and layer A were negatively correlated, suggesting that inputs dominated by the MIN and layer A were concentrated in independent sets of patches. These results indicate that the anatomically observed patchy projections reflect the functional consequences of geniculate lamination. The A layers are high-acuity relays, whereas the MIN is probably a specialization for dim-light vision (Lee et al., 1984; Lee et al., 1992). We propose that the partial overlap of inputs dominated by the A layers and the MIN allows dynamic shifts in their relative contributions to LS responses, optimizing the balance of high-acuity and high-sensitivity channels over a wide range of illumination conditions.

Neuronal responses in extrastriate cortex to objects in optic flow fields.

During locomotion, observers respond to objects in the environment that may represent obstacles to avoid or landmarks for navigation. Although much is known about how visual cortical neurons respond to stimulus objects moving against a blank background, nothing is known about their responses when objects are embedded in optic flow fields (the patterns of motion seen during locomotion). We recorded from cells in the lateral suprasylvian visual area (LS) of the cat, an area probably analogous to area MT. In our first experiments, optic flow simulations mimicked the view of a cat trotting across a plain covered with small balls; a black bar lying on the balls served as a target object. In subsequent experiments, optic flow simulations were composed of natural elements, with target objects representing bushes, rocks, and variants of these. Cells did not respond to the target bar in the presence of optic flow backgrounds, although they did respond to it in the absence of a background. However, 273/423 cells responded to at least one of the taller, naturalistic objects embedded in optic flow simulations. These responses might represent a form of image segmentation, in that cells detected objects against a complex background. Surprisingly, the responsiveness of cells to objects in optic flow fields was not correlated with preferred direction as measured with a moving bar or whole-field texture. Because the direction of object motion was determined solely by receptive-field location, it often differed considerably from a cell's preferred direction. About a quarter of the cells responded well to objects in optic flow movies but more weakly or not at all to bars moving in the same direction as the object, suggesting that the optic flow background modified or suppressed direction selectivity.

A metabolic mapping study of orientation discrimination and detection tasks in the cat.

Increasing evidence suggests that a large number of distinct cortical areas and associated subcortical structures participate in the processing of visual information and that different aspects of visual scenes are evaluated in different areas. This necessitates identification of cortical and subcortical regions cooperating in particular visual tasks. Using the 2-deoxyglucose technique, we monitored the differential activation of areas in the cat visual cortex participating in an orientation discrimination and a detection task. Concordant with previous lesion studies, we found increased activity levels in area 17 in the discrimination condition relative to the detection condition. In addition, the 2-deoxyglucose technique revealed discrimination-related increased activations in the claustrum, the putamen and in parts of the anteromedial, anterolateral and posterolateral lateral suprasylvian visual areas. Regions activated differentially with the detection task comprised subdivisions of areas 17, 18, 19 and 21, posterior area 7 (7p), several areas of the posterior part of the middle and posterior suprasylvian sulcus, the pulvinar complex and the superior colliculus. These results show that the 2-deoxyglucose technique is useful to investigate cognitive brain functions, and that different sets of cortical and subcortical regions are activated during two visual tasks with similar visual stimulation.

Excitatory convergence of Y and non-Y information channels on single neurons in the PMLS area, a motion area of the cat visual cortex.

We analysed the receptive field properties of neurons in the posteromedial lateral suprasylvian (PMLS) visual cortical area of anaesthetized cats in which there was selective conduction block of the largest (Y-type) fibres in one optic nerve. As in normal cats, in cats with selective block of one optic nerve the great majority of PMLS cells could be activated by photic stimulation through either eye. However, the responses evoked by stimulation via the eye with the selectively pressure-blocked optic nerve ('Y-blocked eye') were significantly weaker than those of the same cells evoked by the stimulation via the normal eye. Accordingly, eye dominance histograms were shifted markedly in favour of the normal eye. Furthermore, there was a significant shift towards lower preferred velocities when PMLS cells were photically stimulated via the Y-blocked eye. Finally, when stimulated via the Y-blocked eye, PMLS cells responded poorly or not at all to high stimulus velocities (> or = 100 degrees/s). On the other hand, a number of receptive field properties, such as the spatial organization of receptive fields, the size of the discharge fields, orientation tuning and direction selectivity indices, were not significantly affected by the removal of the Y input. We conclude that virtually all neurons in the PMLS area of the cat receive excitatory input from both Y and non-Y information channels, although the Y channel provides the dominant input and makes the principal contribution to the detection of high-velocity motion.

Simulated optic flow and extrastriate cortex. I. Optic flow versus texture.

A locomoting observer sees a very different visual scene than an observer at rest: images throughout the visual field accelerate and expand, and they follow approximately radial outward paths from a single origin. This so-called optic flow field is presumably used for visual guidance, and it has been suggested that particular areas of visual cortex are specialized for the analysis of optic flow. In the cat, the lateral suprasylvian visual area (LS) is a likely candidate. To test the hypothesis that LS is specialized for analysis of optic flow fields, we recorded cell responses to optic flow displays. Stimulus movies simulated the experience of a cat trotting slowly across an endless plain covered with small balls. In different simulations we varied the size of balls, their organization (randomly or regularly dispersed), and their color (all one gray level, or multiple shades of gray). For each optic flow movie, a "texture" movie composed of the same elements but lacking optic flow cues was tested. In anesthetized cats, > 500 neurons in LS were studied with a variety of movies. Most (70%) of 454 visually responsive cells responded to optic flow movies. Visually responsive cells generally preferred optic flow to texture movies (69% of those responsive to any movie). The direction in which a movie was shown (forward or reverse) was also an important factor. Most cells (68%) strongly preferred forward motion, which corresponded to visual experience during locomotion.

Simulated optic flow and extrastriate cortex. II. Responses to bar versus large-field stimuli.

In the preceding paper we described the responses of cells in the cat's lateral suprasylvian visual area (LS) to large-field optic flow and texture movies. To assess response properties such as direction selectivity, cells were also tested with moving bar stimuli. We expected that there would be good agreement between response properties elicited with optic flow movies and those revealed with bar stimuli. We first asked how well bar response properties predicted responsiveness to optic flow movies. There was no correlation between responsiveness to movies and the degree of end-stopping, length summation, or preference for bars that accelerated and expanded. We then considered only the 322 cells that responded to both bars and optic flow or texture movies and asked how well the strength of their response to movies could be predicted from the direction-tuning curves generated with bar stimuli. One-third of these cells responded much more strongly to movies than could be predicted from their direction-tuning curves. Generally, such cells were rather well tuned for the direction of bar motion and preferred a direction substantially different from what they saw in optic flow movies. Optic flow movies shown in the forward direction were the most effective variety of movie for two-thirds of these cells. To see whether this outcome stemmed from differential direction tuning for bars and large multielement displays, in a second series of experiments we compared direction tuning for bars and large-field texture movies. Many cells showed substantially different direction tuning for the two kinds of stimulus: almost 1/3 of 409 cells had tuning curves that overlapped each other by < 50%. But only a small number of cells (< 10%) responded much better to texture movies than to bars in the predominant direction of image motion in optic flow movies. This result, like that reported in the preceding paper, suggests that cells in LS respond differently to optic flow than to texture displays lacking optic flow motion cues.

Visual motion processing in the anterior ectosylvian sulcus of the cat.

1. Neurons that are selectively sensitive to the direction of motion of elongated contours have been found in several cortical areas in many species. However, in the striate cortex of the cat and monkey, and the extrastriate posteromedial lateral suprasylvian visual area of the cat, such cells are generally component motion selective, signaling only the direction of movement orthogonal to the preferred orientation; a direction that is not necessarily the same as the motion of the entire pattern or texture of which the cell's preferred contour is part. The primate extrastriate middle temporal area is the only cortical region currently known to contain a substantial population of pattern-motion-selective cells that respond to the shared vector of motion of mixtures of contours. 2. From analyzing published data on the connectivity of the cat's cortex, we predicted that the anterior ectosylvian visual area (AEV), situated within the anterior ectosylvian sulcus, might be a higher-order motion processing area and thus likely to contain pattern-motion-selective neurons. This paper presents the results of a study on neuronal responses in AEV. 3. Ninety percent of AEV cells that responded strongly to drifting grating and/or plaid stimuli were directionally selective (directionality index > 0.5). For this group, the mean directionality index was 0.75. Moreover, 55% of these cells were unequivocally classified as pattern motion selective and only one neuron was classified as definitely component motion selective. Thus high-level pattern motion coding occurs in the cat extrastriate cortex and is not limited to the primate middle temporal area. 4. AEV contains a heterogeneous population of directionally selective cells. There was no clear relation between the degree of directional selectivity for plaids or gratings and the degree of selectivity for pattern motion or component motion. Nevertheless, 28% of the highly responsive cells were both more strongly modulated by plaids than gratings and more pattern motion selective than component motion selective. Such cells could correspond to a population of "selection units" signaling the salience of local motion information. 5. AEV lacks global retinotopic order but the preferred direction of motion of neurons (rather than axis of motion, as in the middle temporal area and the posteromedial lateral suprasylvian visual area) is mapped systematically across the cortex. Our data are compatible with AEV being a nonretinotopic, feature-mapped area in which cells representing similar parts of "motion space" are brought together on the cortical sheet.

Chapter 17 Extrinsic and intrinsic connections of the cat's lateral suprasylvian visual area

The lateral suprasylvian visual area (LS) is known to have numerous interconnections with visual cortical areas as well as with subcortical structures implicated in visually-guided behaviors. In contrast, little data is available regarding connections within the LS itself. In order to obtain information about intra-areal connections and to re-investigate LS connectivity with various cortical and subcortical areas, the traces (biocytin or WGA-HRP) was injected into various loci along the medial and lateral banks of the LS. The anterograde tracer, biocytin injections into both medial and lateral bank produced label contained within the respective bank that extended rostrally and caudally from the infection site. In addition, following medical bank injections, considerable label was distributed throughout the fundus and, to a lesser extent, in the lateral bank. In contrast, no label could be detected in the medial bank after lateral bank injections, and, although label was observed in the fundus, it was restricted to the most lateral aspects. Moderate labeling could be observed in the medial bank following the tracer injection into the most rostral aspect of the lateral bank. It is likely that input derived from various visual cortical areas which project to the medial bank of the LS has access to this intra-areal circuitry. This may provide a route by which visual cortical information can be relayed to other cortical and subcortical structures involved in visually-guided behaviors such as the anterior ectosylvian visual cortex, striatum, and the deep layers of the superior colliculus, despite the fact that these structures themselves do not receive substantial direct projections from the visual cortical areas that are associated with the medial bank. Examination of the laminar location of the cells-of-origin of striate and extrastriate projections to LS using retrograde trace, WGA-HRP, revealed that the supragranular laminae of areas 17, 18 and 19 were the source of LS afferents whereas afferents from the other cortical areas (e.g., 20a, 20b, 21a, 21b, 7 and anterior ectosylvian visual area) were from both supra- and infragranular laminae. In addition, all LS subregions received intra-areal afferent projections from all LS cortical laminae. Thus, although rather clear hierarchical relationship between LS and visual cortical areas appears to exist, the interconnections among LS subregions provide no clear evidence of simple hierarchical relationships between regions LS or may have feed-forward and feed-back pathways.

Preservation of Pointing Accuracy Toward Moving Targets After Extensive Visual Cortical Ablations in Cats

Impairments in reaching toward stationary and moving targets were studied in cats after restricted or extensive removal of visual cortical areas (areas 17, 18 and 19 and lateral suprasylvian visual areas). Regardless of the extent of the cortical lesion, cats were at first unable to localise and reach for a stationary target whereas they were soon able to detect and accurately point toward a mobile one. Moreover, the onset latency of such movements was dramatically increased. During post-operative re-training, the cats were unable to improve their accuracy scores when reaching towards stationary targets. In contrast, full compensation was observed for the accuracy of reaching movements directed toward moving targets. A partial recovery was observed for movement latency values that progressively decreased but left a permanent 30–40 ms impairment following extensive lesions. The role of extrageniculate messages and alternative routes involving other cortical areas in taking in charge the visuomotor activity is discussed.

Transient expression of calbindin-D28k immunoreactivity in layer V pyramidal neurons during postnatal development of kitten cortical areas

Calbindin-D28k is a 28 kDa calcium binding protein that has been shown to colocalize with a specific subpopulation of γ-aminobutyric acid inhibitory interneurons in mammalian neocortex. We have examined the ontogeny of calbindin in neonatal kitten cortex in areas 17,18,19,7, medial and lateral suprasylvian visual areas, splenial visual area and cingulate cortex from the day of birth (P0) through maturation of the brain (P101). Transient staining of immature layer V pyramidal cells was seen in kittens six weeks old and younger. This transient staining of pyramidal cells was most intense and the stained neurons were most numerous in cingulate cortex. Apical dendrites of pyramidal cells in cingulate cortex were prominently stained and could be followed to layer I, where they were seen to branch extensively. There were very few calbindin immunoreactive pyramidal cells in primary cortical areas postnatally. Transient staining in extrastriate visual cortical areas disappeared first from the lateral suprasylvian areas, and persisted longest in area 7. Pyramidal neurons in the cingulate gyrus expressed calbindin longest, but calbindin expression by pyramidal neurons ceased by the sixth postnatal week in all areas of the brain.