Extrastriate Cortical Areas Research Articles

Synchronization of Neuronal Activity during Stimulus Expectation in a Direction Discrimination Task

The dorsal pathway of the primate brain, especially the middle temporal area (MT or V5) and the superior middle temporal area (MST or V5a), is strongly involved in motion detection. The relation between neural firing rates and psychophysical performance has led to the assumption that the neural code used by these areas consists of the relative discharge rates of neuronal populations. As an additional neural code, temporal correlation of neural activity has been suggested. Our study addresses the involvement of such a code in awake monkeys performing a motion discrimination task. We found significant temporal correlations between simultaneously recorded pairs of units in areas MT and MST and other extrastriate cortical areas. Units recorded from the same electrode were more frequently synchronized than units recorded from different electrodes placed within the same or different cortical areas. Activity synchronization was present in the expectation period before stimulus presentation and could not be induced de novo by the stimulus. Rather, we found a contrast-dependent reduction of correlation strength on stimulus onset. Correlation strength did not vary systematically with stimulus directions. We conclude that under the conditions of this study, temporal decorrelation of MT and MST neurons could be used to detect the stimulus, but synchronization does not convey specific information about its direction of motion and therefore is unlikely to contribute to performance in our direction discrimination task. Activity synchronization in the period before stimulus onset could be related to attentive expectation.

Contribution of area 17 to cell responses in the striate-recipient zone of the cat's lateral posterior-pulvinar complex.

The cat's lateral posterior-pulvinar complex (LP-pulvinar) contains three main representations of the visual field. The lateral part of the LP nucleus (LPl or striate-recipient zone) is the only region of these extrageniculate nuclei which receives afferents from the primary visual cortex. We investigated the contribution of area 17 to the response properties (orientation and spatial frequency tuning functions) of LPl neurons by cooling or lesioning the visual cortex. Responses of 40 LPl cells were studied before, during and after the reversible cooling of the striate cortex. When tested for orientation, a total of 10 units out of 28 was affected (36%). For most of these cells (eight of 10), cooling the visual cortex yielded a reduction of the cells' visual responses without altering their orientation-selectivity (there was no significant change in the orientation tuning width). For only two cells, inactivation led to an increase in the response amplitude. Also, blocking the visual cortex never modified the direction-selectivity of LPl cells. When tested for spatial frequency, 12 neurons out of 33 were affected (36%) by the experimental protocol. In most cases, we observed a reduction in the responses at each spatial frequency tested, with no change in tuning bandwidth. For only three LPl cells, the effects of inactivation of the visual cortex were restricted to specific spatial frequencies, altering the profile of the spatial frequency tuning function. In five cats, removing area 17 reduced the proportion of visual neurons in LPl and the spared visually evoked responses were noticeably depressed. Despite the reduction in responsiveness, a few LPl receptive fields within the cortical scotoma were still sensitive to the orientation and/or direction of a moving stimulus. This last observation suggests that some properties in LPl could be generated either by circuits intrinsic to the LPl or by afferents from extrastriate cortical areas. Overall, these results indicate that projections from the visual cortex to the striate-recipient zone of the LP-pulvinar complex are mainly excitatory. Despite the strong impact of the area 17 projections, our data suggest that the extrastriate cortex could also play a role in the establishment of response properties in the cat's LPl.

Blindsight in man and monkey

In man and monkey, absolute cortical blindness is caused by destruction of the optic radiations and/or the primary visual cortex. It is characterized by an absence of any conscious vision, but stimuli presented inside its borders may nevertheless be processed. This unconscious vision includes neuroendocrine, reflexive, indirect and forced-choice responses which are mediated by the visual subsystems that escape the direct cerebral damage and the Ensuring degeneration. While extrastriate cortical areas participate in the mediation of the forced-choice responses, a concomitant striate cortical activation does not seem to be necessary for blindsight. Whether the loss of phenomenal vision is a necessary consequence of striate cortical destruction and whether this structure is indispensable for conscious sight are much debated questions which need to be tackled experimentally.

Limits of parallel processing: excitatory convergence of different information channels on single neurons in striate and extrastriate visual cortices.

1. It has been postulated that the distinct parallel retino-geniculo-cortical information channels characterizing visual pathways of virtually all mammals are selectively linked to parallel motion, colour and/or form information processing 'streams' distinguishable within the primary visual cortices, extrastriate cortical areas of occipital lobes and the temporal and parietal visual cortices. 2. Using selective pressure-blocking of the large-fibre channel (the so-called Y-channel) in the optic nerve of the cat, we have experimentally examined the 'selective excitatory parallel links' hypothesis. We conclude that the majority of neurons in the primary visual cortices (areas 17, 18) as well as in the two 'higher order' visual areas, area 21a and posteromedial lateral suprasylvian (PMLS) area, constituting, respectively, part of the 'form' and part of the 'motion' processing streams, receive their excitatory inputs from both Y- and non-Y-information channels. In areas 17, 18 and 21a (but not in PMLS area), there are, however, subpopulations of cells that apparently receive excitatory inputs from only one information channel. 3. Review of the relevant work on the macaque monkey suggests that the situation is similar in the primate: that is, there is a substantial degree of excitatory convergence of different retino-geniculo-cortical information channels on single neurons in the primary visual cortices and the extrastriate cortices constituting parts of the form/colour or the motion processing streams. 4. Despite this high degree of excitatory convergence of different information channels, the large-fibre channels (the Y-channel in the cat and the magnocellular or Y-like channel in macaque), are in both carnivores and primates the principal contributors to the motion processing cortical streams.

Binocular interactions and disparity coding in area 21a of cat extrastriate visual cortex.

We have examined, using both qualitative and quantitative techniques, binocular interactions of extracellularly recorded single neurons in the extrastriate cortical area 21a of anaesthetized and paralysed cats. Consistent with previous reports we have found that: (a) all area 21a neurons were orientation-selective, with about 65% of them preferring orientations within 30 degrees of the vertical; and (b) over 75% of area 21a cells could be activated through either eye. Furthermore, a significant minority (4 cells; about 10%) of a subpopulation of 39 neurons in which binocular interactions were examined quantitatively, were "obligatory binocular neurons", that is, they responded very weakly, if at all, to the monocular stimuli presented through either eye but responded vigorously to simultaneous stimulation through both eyes. Almost 70% (27/39) of neurons tested quantitatively for binocular interaction have shown significant modulation (over 50%) of their peak responses in relation to binocular positional retinal disparities. The majority of neurons sensitive to binocular positional disparities resembled either "tuned excitatory" (22 cells; 56.5% of the sample) or "tuned inhibitory" (2 cells; 5% of our sample) cells. In particular, they gave, respectively, maximal or minimal responses to optimally oriented, moving photic stimuli when the receptive fields plotted through each eye completely or partially overlapped. Although neurons recorded in area 21a have relatively large receptive fields (mean width 3.3 +/- 1.1 degrees; range 2.0-5.6 degrees), the mean width of the disparity tuning curve (2.8 +/- 1.0 degrees; range 1.3-4.8 degrees) for our sample of area 21a neurons was similar to those of neurons with significantly smaller receptive fields, recorded in areas 17 and 18 of cat's primary visual cortex. We conclude that area 21a of the cat, like areas 17 and 18 of primary visual cortex, is likely to play an important role in binocular depth discrimination and might constitute a "higher order" area for stereoscopic binocular vision.

Orientation discrimination in the cat: its cortical locus II. Extrastriate cortical areas.

Luminance-defined edges or bars are among the basic units of visual analysis: a "primitive" component of perception. We have utilized this stimulus in a psychophysical study of bar orientation discrimination in the cat before and after selective lesions in visual cortical areas. The cortices have been divided on the basis of their connectivity into three tiers. Tier I refers to areas 17 and 18, tier II includes areas that receive directly from tier I, and tier III includes those areas that receive directly from tier II. Previous studies (Vandenbussche et al. [1991] J. Comp. Neurol. 305:632-658) have shown that the discrimination of bar orientation depends heavily upon the integrity of areas 17 and 18 (tier I). The present study indicates that several extrastriate areas in tiers II and III contribute to this discrimination task. Our data suggest that the anterior medial lateral suprasylvian, the posterior lateral lateral suprasylvian (tier II), and the anterior lateral lateral suprasylvian (tier III) areas are most likely to contribute to bar orientation discrimination.

Orientation discrimination in the cat: Its cortical locus II. Extrastriate cortical areas

Orientation discrimination in the cat: Its cortical locus II. Extrastriate cortical areas

Induction of c- fos protein by patterned visual stimulation in central visual pathways of the rat

Localized patterned visual stimulation was used in rats to investigate the feasibility of stimulus-dependent induction of the immediate early gene c- fos in neurons of cortical and subcortical visual centers of this mammal. Moving and stationary visual patterns, consisting of gratings and arrays of dark dots, induced Fos-like immunoreactivity in populations of neurons in retinotopically corresponding stimulated regions of the dorsal and ventral lateral geniculate nucleus (dLGN, vLGN), stratum griseum superficiale of the superior colliculus, nucleus of the optic tract, and primary (striate) visual cortex. Only moving stimuli induced Fos-like immunoreactive (FLI) neurons in extrastriate visual areas, particularly in the anterolateral (AL) visual area. This suggests that area AL is equivalent to the motion sensitive areas MT and PMLS of the monkey and cat. Stimulus-induced FLI neurons in the striate cortex were predominantly distributed in layers 4 and 6, while few labeled neurons were present in layers 2–3, and almost none in layer 5. The laminar distribution of stimulus-induced FLI cells in the extrastriate cortical area AL was similar to that of the striate cortex, with the exception that more FLI cells were present in layer 5. Statistical comparison of somata size of the stimulus-induced FLI neurons in dLGN with that of Cresyl violet stained neurons in the same sections revealed that the population of geniculate FLI neurons is composed of relay cells and interneurons.

Borders of multiple visual areas in humans revealed by functional magnetic resonance imaging.

The borders of human visual areas V1, V2, VP, V3, and V4 were precisely and noninvasively determined. Functional magnetic resonance images were recorded during phase-encoded retinal stimulation. This volume data set was then sampled with a cortical surface reconstruction, making it possible to calculate the local visual field sign (mirror image versus non-mirror image representation). This method automatically and objectively outlines area borders because adjacent areas often have the opposite field sign. Cortical magnification factor curves for striate and extrastriate cortical areas were determined, which showed that human visual areas have a greater emphasis on the center-of-gaze than their counterparts in monkeys. Retinotopically organized visual areas in humans extend anteriorly to overlap several areas previously shown to be activated by written words.

Visual latencies in cytochrome oxidase bands of macaque area V2.

Cytochrome oxidase bands in area V2 of the primate visual cortex constitute separate relays for parallel channels relaying information from area V1 to other extrastriate cortical areas. We investigated whether information is transferred at the same speed in the different channels by measuring the latencies of neurons in different cytochrome oxidase bands identified by the presence or absence of retrogradely labeled cells from injections in area V4. Results show that neurons in the thick and pale bands respond 20 msec earlier than those in the thin bands. We also found that color-selective neurons respond later than neurons with no selectivity for color and that direction-selective neurons have shorter latencies than neurons with no selectivity for the direction of stimulus movement.

The development of parvalbumin and calbindin-D28k immunoreactive interneurons in kitten visual cortical areas

Calbindin-D and parvalbumin are calcium binding proteins which are found in non-overlapping subpopulations of GABA-ergic interneurons in mammalian neocortex. We studied the development of these calcium-binding proteins in interneurons of cat striate and extrastriate cortical areas which have differing patterns of connectivity and follow different developmental timetables. We examined primary visual areas 17 and 18, secondary visual area 19, medial lateral suprasylvian and lateral suprasylvian areas (MLS and LLS) and association areas 7 and the splenial visual area from the day of birth (P0) through P101. Parvalbumin-immunoreactive (ir) interneurons followed the inside-out pattern of maturation of cortical laminae. They were located only in infragranular layers at the earliest ages and were not observed in the overlying cortical plate. At 3 weeks of age, when cortical lamination is mature, parvalbumin stained cells were found in all cortical layers except layer I. The number of stained secondary and tertiary dendrites in the parvalbumin-ir interneuronal population decreased with age. This change was associated with a shift in the molecular weight of parvalbumin detected on Western blots. During the first postnatal week, the area 17 18 border contained more parvalbumin-ir neurons than other visual areas. The developmental pattern of calbindin staining differed considerably from the parvalbumin staining pattern. Very few calbindin-ir interneurons were seen in area 17 during the first 2 weeks of life. In lateral cortical areas, calbindin-ir neurons were located in cortical plate, infragranular layers of cortex and white matter/subplate. Calbindin-ir neurons increased in supragranular layers of secondary cortical areas by P7 and in area 17 by P20. In the mature cortex, the calbindin staining pattern was bilaminar, with a dense band of calbindin-ir cells in layer II and a second band in layers V–VI. There was no difference in the distribution of calbindin-ir neurons among visual areas at maturity.

Retinotopy of cortical connections between the striate cortex and extrastriate visual areas in the rat.

Previous studies have determined that the striate cortex of the rat is reciprocally connected with multiple extrastriate cortical areas that are retinotopically organized. The objective of this study was to investigate the retinotopy of the striate-extrastriate connections in the rat, by placing triple or double injections of fluorescent tracers (fluorogold, fast blue, rhodamine dextran, or rhodamine-labeled microspheres) in different regions of the striate cortex (Oc1) and mapping the distribution of cells and fibers labeled with the different tracers in the lateral (Oc2L) and medial (Oc2M) extrastriate cortex. The tracer injection sites were visualized in tangential sections of the flattened cortex and correlated with the myelin layout of the striate cortex and with an electrophysiological map from previous studies. The results showed retinotopically organized Oc1 connections with ten different extrastriate cortical areas. The location of these extrastriate areas and the retinotopy of their striate connections remained mostly invariant despite changes of the injection sites in Oc1. Thus, the quadrantic retinotopy was obtained for striate connections to areas posterior, posterolateral, lateromedial, laterointermediate, laterolateral, anterolateral and rostrolateral in Oc2L; and to areas posteromedial, anteromedial, and anterior in Oc2M. The present anatomical map correlates well with electrophysiological maps of the rat extrastriate cortex from previous studies. Furthermore, they provide a definition of the retinotopy of some areas that have not been completely mapped before. These results reaffirm the existence of multiple extrastriate visual areas in the rat.

Visual activity in areas V3a and V3 during reversible inactivation of area V1 in the macaque monkey

1. Behavioral studies in the monkey and clinical studies in humans show that some visuomotor functions are spared in case of a V1 lesion. This residual vision appears to be subserved at least partially by visual activity in extrastriate cortex. Earlier studies have demonstrated that neurons in area V2 lose their visual responses when V1 is reversibly inactivated. On the other hand, Rodman and collaborators have recently shown that neurons in the middle temporal area (area MT) remain visually responsive when V1 is lesioned or inactivated. The purpose of the present study was to determine whether area MT is unique among extrastriate cortical areas in containing visually responsive neurons in the absence of input from area 17. 2. A circular part of the opercular region of area V1 was reversibly inactivated by cooling with a Peltier device. In that condition, 149 sites were recorded in the retinotopically corresponding regions of areas V3 and V3a. 3. About 30% of sites in area V3a still responded to visual stimulation when V1 was inactivated. On the contrary, nearly all sites in area V3 ceased to fire to visual stimulation. Receptive-field properties were assessed with qualitative measures; for most single cells or multiunit sites that responded during V1 inactivation, these properties did not change during cooling. 4. These results suggest that area V3a could take part in spared visuomotor abilities in case of a lesion of V1. Areas V3a and MT are both part of the occipitoparietal pathway, which suggests that the residual vision observed after a lesion of area 17 may depend mostly on this pathway.

Disjunctive eye movement evoked by microstimulation in an extrastriate cortical area of the cat

Slow disjunctive eye movement similar to ocular convergence was evoked by microstimulation in parts of the lateral suprasylvian area (LSA) in alert cats. A tungsten-in-glass microelectrode was used for stimulation, and eye movement was monitored using the magnetic search coil method. The velocity-versus-amplitude relationship of disjunctive eye movement evoked by microstimulation was comparable to that of ocular convergence evoked by presenting a visual target. It is suggested that the LSA plays a role in controlling convergence eye movement.

Visual corticopontine and tectopontine projections in the macaque.

We studied projections from extrastriate visual areas and the superior colliculus to the pontine nuclei of monkeys using degeneration staining and transport of wheatgerm agglutinin horseradish peroxidase, and 3H amino acids. The superior colliculus and the extrastriate cortical visual areas both project to the ipsilateral dorsolateral region of the pontine nuclei. The projections from extrastriate visual cortex occupy a much larger territory within the pontine nuclei than those from the superior colliculus. The superficial laminae of the superior colliculus project only to the ipsilateral pontine nuclei. The projection to the contralateral nucleus reticularis tegmenti pontis arises from cells in deeper laminae within the superior colliculus.

Corticocortical connections of anatomically and physiologically defined subdivisions within the inferior parietal lobule

The anatomical and functional organization of the inferior parietal lobule was investigated in macaque monkeys by using anterograde and retrograde anatomical tracing techniques and single cell recording techniques in awake, behaving monkeys. The connections of areas 7a and 7b, and of two previously unexplored areas, the lateral intraparietal area (LIP) and the dorsal prelunate area (DP), were examined in detail. Functional mapping experiments were performed in all four areas. Prior to this study the pathways for visual input to area 7a were unclear. In these experiments we found several direct projections from extrastriate visual areas, including the lateral intraparietal (LIP), dorsal prelunate (DP), parieto-occipital (PO), and medial superior temporal (MST) areas into area 7a. Using the observed laminar patterns of connections between areas 7a, LIP, and DP and other extrastriate cortical areas, we were able to construct a hypothetical flow of visual information processing from striate cortex to area 7a. A broader hierarchy was also produced, which relates the positions of areas 7a, 7b, LIP, and DP to various cortical fields in the parietal, temporal, and frontal lobes. By combining single cell recording techniques in trained monkeys with anatomical tracing techniques, we have parceled the inferior parietal lobule into several subdivisions on the basis of both anatomical and physiological grounds. A clear segregation of visual and somatosensory responses was found in the inferior parietal lobule with areas 7a, LIP, and DP being visual and visual-motor and area 7b being primarily somatosensory. A similar segregation was found anatomically with areas 7a, LIP, and DP being interconnected primarily with other visual cortical areas and area 7b being connected with several somatosensory areas. Area 7b was also found to connect to a few visual cortical areas, and these connections likely account for the small but consistent number of visually responsive cells that are found in this region. Areas LIP, DP, and 7a differed in receptive field and saccade-related properties. Area 7a visual receptive fields were very large and usually bilateral with a small but significant number of them having receptive field centers in the ipsilateral visual field. Area DP and LIP receptive fields were smaller and the receptive field peaks were almost always confined to the contralateral visual field. Areas 7a, DP, and LIP all contained cells with saccade-related responses; however, in area 7a there were fewer saccade cells than area LIP, and presaccadic responses were only observed in area LIP.(ABSTRACT TRUNCATED AT 400 WORDS)

Visual response properties of cortical inputs to an extrastriate cortical area in the cat

The existence of multiple areas of extrastriate visual cortex raises the question of how the response properties of each area are derived from its visual input. This question was investigated for one such area in the cat, referred to here as the Clare-Bishop area (Hubel & Wiesel, 1969); it is the region of lateral suprasylvian cortex that receives input from area 17. A novel approach was used, in which kainic acid was injected locally into the Clare-Bishop area, making it possible to record directly from afferent inputs. The response properties of the great majority of a sample of 424 presumed afferents resembled cells in areas 17 and 18. Thus, a systematic comparison was made with cells from area 17's upper layers, the source of its projection to the Clare-Bishop area (Gilbert & Kelly, 1975), to see whether these afferents had distinctive properties that might distinguish them from cells projecting to areas 18 or 19. Some differences did emerge: (1) The smallest receptive fields typical of area 17 were relatively scarce among afferents. (2) Direction-selective afferents were more abundant than were such cells in area 17. (3) End-stopped afferents were extremely rare, although end-stopped cells were common in area 17's upper layers. Despite these differences, afferents were far more similar in their properties to cells in areas 17 and 18 than to cells in the Clare-Bishop area. Compared to the latter, afferents showed major discrepancies in receptive-field size, in direction selectivity, in end-stopping, and in ocular dominance distribution. These differences seem most likely to stem from circuitry intrinsic to the Clare-Bishop area.

Chapter 4: Superior colliculus-mediated visual behaviors in cat and the concept of two corticotectal systems

This chapter is the outgrowth of a series of experiments in cat in an attempt to understand how cortex and midbrain (specifically, the superior colliculus) interact in the production of visual behaviors. A number of years ago, researchers examined how primary visual cortex affects the most visually “dominant” area of the superior colliculus (SC), its superficial laminae. This proved to be an excellent model for determining how the brain builds receptive fields from converging afferents. An extrastriate cortical area has now been identified as controlling the deep laminae SC cells that connect to those areas of the brainstem and spinal cord through which overt responses are initiated. Compromising any station along this circuit seriously impairs visual attention and orientation capability. Many of the receptive field properties of these visual cells were found to be dependent on the inputs received from primary visual cortex. This dependency is apparent when removal of striate cortex eliminated from these SC cells their binocularity, directional selectivity, and preferences for moving rather than stationary flashed.

Topography of the afferent connectivity of area 17 in the macaque monkey: a double-labelling study.

The various structures afferent to area 17 (or V1) of the macaque monkey have widely differing retinotopic organizations. It is likely that these differences are reflected in the topographic organizations of the projections from these structures to area V1. We have investigated this issue by placing side-by-side injections of two retrograde fluorescent tracers, fast blue and diamidino yellow, in V1. By examining the extent of mixing of the two populations of singly labelled cells and the presence of doubly labelled cells, in different structures, we have characterized the topography of each projection in terms of the size of its axonal arborization and the amount of convergence and divergence. The afferents from the lateral geniculate nucleus (LGN) and from the pulvinar are organized in a point-to-point fashion. The maximum extent of axonal arborization of these afferents is 0.5 mm and these projections demonstrate little scatter (i.e., neighboring LGN neurons project to adjacent regions of V1). The other two subcortical structures examined, the claustrum and the intralaminar nuclei, demonstrate a much larger scatter and wider axonal arborizations in their projections to V1 than do the LGN and pulvinar. Two-dimensional reconstructions were made of the distribution of labelled neurons in extrastriate cortical areas. Using the separation between patches of labelled cells and transitions in myelin-stained sections, we have identified seven separate cortical regions containing labelled cells. Two of these can be identified as area V2 and the middle temporal visual area (MT). Three other regions correspond to areas V3, V3A and V4t. Finally, two more regions of labelling have been distinguished that belong to area V4. These results demonstrate that, at least within the central 6 degrees of visual field, all the presently known extrastriate visual cortical areas project to V1. This result is interesting in view of the fact that only a few extrastriate cortical areas are reported to receive afferents from V1. Three groups of cortical areas can be distinguished on the basis of the characteristics of their cortical connections to V1. The first group contains area V2, V3, and the posterior region of V4. These areas project to V1 with infra- as well as supragranular layer neurons and show limited axonal arborization and scatter in the projection. The second group consists of two regions of labelling in the superior temporal sulcus corresponding to V4t and MT and another on the annectant gyrus (V3A). These regions contain almost exclusively infragranular labelling and show wide axonal arborization and scatter in their projections to V1.(ABSTRACT TRUNCATED AT 400 WORDS)

Immediate postoperative retention of visual discriminations following selective cortical lesions in the cat

Cats were trained preoperatively for brightness discrimination, and 7 pattern and form discriminations, and then retested for preoperative retention on each discrimination. Cortical lesions were then placed in areas 17 and 18 in one group (4 cats), in areas 17, 18 and 19 in another group (3 cats), and in suprasylvian cortex (areas 7, 21, and parts of 19, 5 and the lateral suprasylvian cortex) in a third group (4 cats). Results are also reported for a fourth group with extensive suprasylvian lesions, to which was added an unintended undercutting of areas 17 and 18 (4 cats). While during original preoperative learning the training continued until a fixed, stringent criterion of performance was attained, both preoperative and postoperative retention was tested in short sessions, involving a limited number of trials and a less stringent statistical criterion (significant run). After extensive removal of areas 17 and 18, all cats behaved as though following the cortical lesion they could immediately recognize the discriminative stimuli as efficiently as before, with no need for retraining. On the contrary, the group with areas 17, 18 and 19 lesions showed a substantial postoperative loss of all discriminations, and especially for the more difficult form discriminations, the reattainment of a significant level of performance was hard or impossible within the allotted number of trials. Also in the group with limited suprasylvian lesions, postoperative retention was generally impaired, but the reacquisition of efficient performance was superior to that of the previous group. Finally, large suprasylvian lesions encroaching on the white matter under areas 17, 18 and 19 proved disruptive for all discriminative capacities, both in retention and in relearning. The excellent retention of all discriminations following areas 17 and 18 lesions once again shows that these areas are by no means essential for complex vision in the cat. In addition, the results strongly indicate that the high-level visual capacities of destriate cats are not due to reorganization of readaptation processes occurring in extrastriate areas after a 17 18 removal. The clear-cut retention deficits which were present in cats with cortical lesions more extensive than areas 17 and 18 or outside of the latter areas prove the essential participation of extrastriate cortical areas in visual discrimination including form. However, the distribution of functions among the various visual cortical areas in visual discrimination remains poorly understood. Attempts at solving this problem will have to consider the variety and the multiplicity of inputs to the array of cortical visual areas as well as the highly interactive nature of the visual system.