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Here we review the results of anatomical and physiological studies in tree shrew visual cortex which focus on the contribution of vertical and horizontal inputs to receptive field center and surround properties of layer 2/3 neurons. A fundamental feature of both sets of connections is the arrangement of axon arbors in a fashion that respects both the orientation preference and retinotopic displacement of the target site. As a result, layer 2/3 neurons receive convergent input from populations of layer 4 and other layer 2/3 neurons whose receptive fields are displaced along an axis in visual space that corresponds to their preferred orientation. Although, horizontal connections extend for greater distances across the cortical surface than vertical connections, the majority of these inputs link neurons with overlapping receptive fields, emphasizing that both feed-forward and recurrent circuits are likely to play a constructive role in generating properties (such as orientation selectivity) that define the receptive field center. Both within and beyond the dimensions of the receptive field center, the distribution of horizontal connections accords remarkably well with the magnitude and axial tuning of length summation effects. Taken together, these results suggest a continuum of functional properties that transcends the traditional designation of receptive field center and surround. By extension, we suggest that the perceptual effects of stimulus context may arise from stimulus interactions within the receptive field center as well as between center and surround. [Copyright &y& Elsevier]

The feedback effect of activity of area 21a on orientation maps of areas 17 and 18 was investigated in cats using intrinsic signal optical imaging. A spatial frequency-dependent decrease in response amplitude of orientation maps to grating stimuli was observed in areas 17 and 18 when area 21a was inactivated by local injection of GABA, or by a lesion induced by liquid nitrogen freezing. The decrease in response amplitude of orientation maps of areas 17 and 18 after the area 21a inactivation paralleled the normal response without the inactivation. Application in area 21a of bicuculline, a GABAa receptor antagonist caused an increase in response amplitude of orientation maps of area 17. The results indicate a positive feedback from high-order visual cortical area 21a to lower-order areas underlying a spatial frequency-dependent mechanism. [Copyright &y& Elsevier]

We present a comprehensive analysis of the intrinsic variability of the periodicity of ocular dominance columns in cat primary visual cortex (area 17) and its relationship to genetic background and visual experience. We characterized the intra-areal and interindividual variability of column spacing in a large set ( n = 49) of ocular dominance patterns adapting a recently developed technique for the two-dimensional analysis of orientation column patterns. Patterns were obtained from three different cat colonies (termed F, M and D), the cats having either normal visual experience or experimentally induced strabismus. Two-dimensional maps of local column spacing were calculated for every pattern. In individual cortices, local column spacings varied by > 50% with the majority of column spacings ranging between 0.6 and 1.5 mm in different animals. In animals from colonies F and M ( n = 29), the mean column spacing ranged between 1.03 and 1.27 mm and exhibited no significant differences, either between the two breeds or between strabismic and normal animals. The mean spacing was moderately clustered in the left and right brain hemisphere of individual animals but not in littermates. In animals from colony D ( n = 2), average column spacing ranged between 0.73 and 0.95 mm, and was thus significantly different from the distribution of spacings in animals from breeds F and M, suggesting an influence of genetic factors on the layout of ocular dominance columns. Local column spacing exhibited a considerable systematic intra-areal variation, with largest spacings along the representation of the horizontal meridian and smallest spacings along the peripheral representation of the vertical meridian. The total variability of ocular dominance column spacing comprised 24% systematic intra-areal variation, 18% interindividual differences of mean column spacing and 58% nonsystematic intra-areal variability. [ABSTRACT FROM AUTHOR]

The primary visual cortex (area 17) of cats continues to grow substantially during early postnatal development. To assess the implications of this growth for the organization of visual cortical maps, we analysed both postnatal growth of area 17 and the spacing of ocular dominance columns in the same animals using 2-deoxyglucose autoradiography to label cortical activation patterns. Quantitative analyses of area size and column spacing were performed on flat-mount sections of the cortical hemispheres. Our analysis of the surface area revealed an average increase of the size of area 17 between the third and tenth postnatal weeks by about 51%. About 75% of this increase occurred during the third and sixth postnatal weeks (size increase of 37%). However, the distance between adjacent columns did not exhibit a similar increase but rather remained constant during the same postnatal period. Since cortical growth is not accompanied by an increased spacing of ocular dominance columns, new functional modules must somehow be added during the first postnatal weeks to occupy the enlarging cortical sheet. Possible mechanisms underlying the formation of new modules are discussed. [ABSTRACT FROM AUTHOR]

During ontogenetic development, the visual cortical circuitry is remodeled by activity-dependent mechanisms of synaptic plasticity. From a dynamical systems perspective this is a process of dynamic pattern formation. The emerging cortical network supports functional activity patterns that are used to guide the further improvement of the network’s structure. In this picture, spontaneous symmetry breaking in the developmental dynamics of the cortical network underlies the emergence of cortical selectivities such as orientation preference. Here universal properties of this process depending only on basic biological symmetries of the cortical network are analyzed. In particular, we discuss the description of the development of orientation preference columns in terms of a dynamics of abstract order parameter fields, connect this description to the theory of Gaussian random fields, and show how the theory of Gaussian random fields can be used to obtain quantitative information on the generation and motion of pinwheels, in the two dimensional pattern of visual cortical orientation columns. [Copyright &y& Elsevier]

In the primary visual cortex of cats, ferrets and macaque monkeys, the thalamocortical afferents conveying signals from the two eyes terminate in alternating regions of layer IV known as ocular dominance columns. Previous experiments have indicated that the periodicity of these columns can be influenced by visual experience: compared to normally raised animals both strabismic cats and cats raised with alternating monocular exposure displayed an increased spacing of adjacent ocular dominance columns in the primary visual cortex (area 17). However, recently it was shown that the formation of ocular dominance columns begins much earlier than previously supposed, indicating that early visual experience might only have a limited influence on the development of the spatial pattern of ocular dominance columns. We therefore visualized the complete pattern of ocular dominance columns in area 17 of normally raised and strabismic kittens during early postnatal development (age 3–6 weeks), particularly focussing on littermates. In addition, we used a previously developed spatial analysis (period statistics) to quantify columnar spacing two-dimensionally. We observed a pronounced interindividual variability in both normally raised and strabismic animals, with column spacings ranging from 783 to 1362 µm. In contrast to previous reports, there were no significant differences in columnar periodicity between normally raised and strabismic cats. These data indicate that rearing has less influence on column spacing while the interindividual variability is much greater than previously supposed, suggesting that genetic differences have an influence on column spacing. [ABSTRACT FROM AUTHOR]

There are two prevailing explanations for the foveal deficit in texture segmentation reported in previous works. One is based on the spatial and temporal properties of the stimuli, which means in terms of physiology a strong contribution of the Magno-channel. The other one is purely spatial and assigns filters of different bandwidths to each eccentricity in the visual field. We have challenged the first explanation experimentally by using isoluminant stimuli. The central performance drop persisted although the Magno-channel is known to respond weakly to stimuli with low luminance contrast. Therefore, we agreed with the spatial explanation. But instead of the abstract filter theories from previous works we propose a computational neural model assuming local lateral interactions in a cortical map model. The psychophysical performance measures could be directly related to geometric properties of the primary visual cortex concerning its mapping geometry and its intrinsic interaction width. Our model accounts quantitatively for our own psychophysical data as well as for others from literature. In general, we claim that the high foveal retino–cortical magnification maps texture elements too far away from each other for being compared by local processes. [Copyright &y& Elsevier]

Elucidation of neural connectivity patterns in the brain are thought to give us a mechanistic understanding of how the brain works. Functional connectivity is best studied by simultaneous recording of single-unit activity from many neurons. Accordingly, various types of multiple-microelectrode systems have been developed. We have studied long-range lateral interactions in cat striate cortex. To physiologically characterize interacting cells recorded simultaneously, we used two microelectrodes whose movements were controlled by two independently-movable microdrives. The tips of the two microelectrodes were separated by ∼2 mm or more. During preliminary plotting of two receptive fields of cell pairs, we often noted the emergence of perfectly synchronous firing between two spike trains (amplitude ratio, about 20:1) registered with two microelectrodes. Synchronously firing, smaller spikes disappeared when larger spikes of the pair were lost to either substantial advancement of or placing an electrolytic lesion at the electrode registering the latter. The synchrony also disappeared when two microdrive systems were shielded individually. We concluded that the synchrony was attained through capacitance coupling between two microdrive systems. We proposed a few practical recommendations to avoid the contamination of cross correlograms with the false-positive, narrow peak at time zero due to the presence of reflected spike trains. [Copyright &y& Elsevier]

Abstract It has been suggested that synchronization of action potentials encodes diverse features of a single image. However, properties of the synchronization, which occurs on a time scale of ≈ 1–5 ms, are still poorly understood. We have tested the modulation of synchronization by manipulating the contextual targets introduced in the surround of the receptive field. Experiments were carried out on anaesthetized cats prepared for multiunit and single-cell recordings in area 17. Initially, a patch of sine-wave drifting grating was positioned over the overlapping receptive fields of several neurons. If this coherent motion produced a significant synchronization in cross-correlograms, contextual targets were added. The first contextual stimuli were two sine-wave patches placed above and below the central compound receptive field. Only the contrast of contextual targets changed. Results show that the larger the differential contrast the higher the synchronization. The second contextual stimulus was a lateral shift of a sine-wave patch. Data show that the wider the distance between the central and peripheral patches the better the synchronization. Furthermore, results suggest that the synchrony pattern computed by cross correlating multiunit recordings from two sites differs when the cross correlation is carried out between individual units belonging to each multiunit recording. Together with our previous results it appears that synchronization is stimulus dependent and its strength increases with larger disparities included in the whole stimulating image. [ABSTRACT FROM AUTHOR]

The aim of this study was to compare three different imaging techniques which are used to provide data on the laminar structure of the human cerebral cortex. Region V1 of Brodmann's area 17 stained with cresyl violet was investigated, and a conventional semi-automatic morphometric evaluation, the videomicroscropic procedure and a new transparent flat-bed scanning technique were compared. The results of each digitizing method were converted into normalized profiles which allow the laminae in the striate cortex to be displayed. It was found that major laminar patterns can be detected by the scanning technique, but that subsidiary laminations are more clearly displayed by morphometry and videomicroscopy. For magnifications up to × 400 a high resolution transparent flat-bed scanner may be used in place of the videomicroscopy technique. [ABSTRACT FROM AUTHOR]

The aim of this investigation was to understand the functions of long horizontal connections projecting from area 17 to area 18 in cats. The animals were anesthetized and prepared for recording single-cell responses to sine-wave gratings in area 18. Neuronal activity was analyzed under three conditions: prior to, during, and after inactivation of a circumscribed region of area 17. The latter was depressed with micro-injections of GABA. Cells in both areas were in close retinotopic correspondence. Cells were classified as simple and complex types. Globally, simple cells were less affected than complex units, and those which were affected shifted their optimal spatial frequency to higher values. Complex neurons were more often influenced by the interruption of area 17 input. Namely, the peaks of the tuning curves were displaced on the x-axis to a new optimal spatial frequency. This effect was obtained by a dual change: a decline in the discharge strength to the optimal spatial frequency and an enhancement to nonoptimal spatial frequency. Contrast sensitivity function disclosed similar shifts of optimal spatial frequencies. Likewise bandwith, spatial resolution, high cutoff, and low cutoff were modified to a greater extent in complex cells. It appears that there is no relationship between areas 17/18 orientation difference and the modifications observed in tuning curves to spatial frequencies. The results suggest that neurons of area 18 may carry multiple-frequency channels and that area 17 facilitates the emergence of one particular spatial frequency. [ABSTRACT FROM AUTHOR]

Receptive field properties of extracellularly recorded units in the visual cortex (area 17) of cats made bilaterally amblyopic by a variety of rearing conditions were measured and compared with the properties of units in normal cats. Properties studied included sensitivity to vernier offset, response facilitation to increasing bar length, receptive field size, responsiveness to moving and flashed stimuli, orientation tuning, the relation between mean firing rate and its variance, the amount of overlap of regions of on and off responsiveness in simple and complex cells, and, for flashed stimuli, latency to response onset, time to peak response, and response decay time constant. Behavioural testing of the amblyopic animals showed that spatial resolution was 2-4 times lower and vernier acuity thresholds 10-20 times greater than normal. Despite this, several neuronal response properties did not differ significantly from those in normal animals. These included peak responsiveness to moving stimuli, widths of orientation tuning curves, response variability, and latency to initial response for flashed stimuli. Other properties showed small but significant changes. Sensitivity to vernier offset (impulses per degree of offset) was reduced to nearly half its normal level; receptive field sizes increased by about 24% and an incomplete segregation of regions of on and off responsiveness was found in some cells, which made them hard to classify as simple or complex. Responses to flashed stimuli were smaller and more persistent. Their statistical significance notwithstanding, it seems unlikely that these relatively small response abnormalities in area 17 can fully account for the observed behavioural deficits. [ABSTRACT FROM AUTHOR]

We examined the orientation tuning curves of 86 cells located in layer V of area 17, before, during, and after focal blockade of a small (300-μm diameter) region of near-retinotopic register in layer V of area 18 of quantitatively established orientation preference. Such focal blockade revealed three distinct populations of area 17 layer V cells-cells with decreased responses to stimuli of some orientations (21%), cells with increased responses to stimuli of some orientations (43%), and cells unaffected by the focal blockade (36%). These effects were clearcut, reproducible, and generally directly related to the known receptive field properties of the cell recorded in area 18 at the center of the zone of blockade. These effects were also analyzed in terms of alterations in orientation bandwidth in the cells in area 17 as a result of the blockade-bandwidth increases (22%) and decreases (24%) were found; however, these changes were essentially unrelated to the measured receptive field properties. Inhibitory and excitatory effects were most pronounced when the regions in areas 17 and 18 were of like ocular dominance and were of similar orientation preference. Inhibitory effects (suggesting a normally excitatory input) were most dependent upon the similarity of receptive fields; excitatory effects (suggesting a normally inhibitory input) were less heavily dependent. [ABSTRACT FROM AUTHOR]

The main input of the lateral part of the cat's lateralis posterior-pulvinar complex (LP-P) comes from the primary visual cortex. We investigated the response properties of cells in area 17 projecting to the striate-recipient zone (LPl) of the cat's LP-P complex. The cells' receptive fields were stimulated with drifting sine-wave gratings. Cells whose fibres terminate in the superior colliculus were also recorded, to determine how their properties compare with those of cortico-LPl cells and to investigate the possibility that LPl is innervated by collaterals of cortico-tectal units. A total of 26 cells in the striate cortex were identified by antidromic activation from the LPl (mean latency 2.2 ms) and 22 from the colliculus (mean latency 2.5 ms). Only six cortical cells could be activated from the LPl and the colliculus. All cortico-LPl cells except for two responded to drifting sinusoidal gratings with unmodulated discharges (AC/DC ratios <1). On the basis of their modulation index, these units were classified as complex cells. All cortico-LPl cells were selective for the orientation of gratings (mean bandwidth of 28°). There was a tendency for cortico-LPl cells to prefer vertical and horizontal orientations. More than half of these cells (57%) were direction selective. Strong orientation anisotropies were also found in the receptive fields of cortico-tectal cells, since almost all units responded preferentially to horizontally oriented gratings. The mean preferred spatial and temporal frequencies of cortico-LPl cells were 0.74 c/deg (bandwidth 2.03 octaves) and 2.7 Hz (bandwidth 2.5 octaves), respectively. These properties did not differ significantly from those of cortico-tectal cells. Most cortico-LPl cells (72%) exhibited contrast-response curves with saturation at low contrast (mean half-saturation 0.2). For the remaining units, the responses increased linearly with contrast without clear saturation. For more than half of cortico-tectal cells (60%), the contrast function was also characterised by a response saturation. Almost all cortico-LPl cells responded to moving random dot patterns with mean tuning functions of 43.6°. 'Standard' as well as 'special' complex cells were found to be equally responsive to the motion of visual noise. Similar properties were recorded for cortico-tectal cells (mean bandwidth of 44.2°). Cortico-LPl and cortico-tectal cells were either binocularly or monocularly driven by the contralateral eye and their mean spontaneous firing rates were 11.7 and 10.9 spikes/s, respectively. These cells were presumably located in layer V. Stimulation of LPl and colliculus also evoked trans-synaptic responses in area 17. The average latency of the orthodromic responses from LPl was much shorter than that from the colliculus (medians 3.5 and 50 ms, respectively). The findings indicate that almost all cortico-LPl units have complex receptive fields and that their overall properties differ from those of recipient cells in LPl. These results also indicate that LPl is not likely to be innervated by collaterals of fibres of cortico-tectal cells. While cortico-LPl and cortico-tectal cells appear to form two distinct populations, there is no significant difference between the overall properties of these two cell groups. [ABSTRACT FROM AUTHOR]

Stimulation of the mesencephalic reticular formation facilitates responses in the visual cortex elicited from the optic radiation. Using intraveneous administration of cholinergic antagonists we investigated in adult cats and two kittens whether this effect is mediated by cholinergic mechanisms. When administered alone the muscarinic antagonists atropine and scopolamine and the nicotinic antagonist mecamylamine failed to block reticular facilitation and sometimes even enhanced the effects of reticular stimulation. However, when administered in combination muscarinic and nicotinic antagonists eliminated or significantly reduced the facilitation. This was even true when the two antagonists were administered with a time lag of several hours. These results support the notion that reticular facilitation of cortical responses is mediated by cholinergic mechanisms and suggest that this effect is mediated either by a receptor with a mixed pharmacological property or by two independent pathways acting via nicotinic and muscarinic receptors. This hypothesis is discussed in the context of recent evidence on cholinergic transmission and earlier data on the pharmacology of reticular arousal. [ABSTRACT FROM AUTHOR]

Cholinergic afferents to the neocortex controlled by the mesencephalic reticular formation (MRF) are known to transiently facilitate cortical excitability. In an attempt to identify the pathway mediating this effect in the cat visual cortex we combined retrograde tracing techniques with immunocytochemical methods to visualize the acetylcholine-synthesizing enzyme choline acetyltransferase (ChAT). In addition we examined, in acute electrophysiological experiments, whether local neurotoxin injections into nuclei of the basal forebrain interfered with the reticular facilitation of cortical evoked potentials. Cholinergic projections to area 17 originate from different centers in the homolateral substantia innominata/internal capsule, the septal nuclei, and the nuclei of the diagonal band of Broca. No direct cholinergic projection from the MRF to the visual cortex was observed. Retrogradely labelled cells intermingled with ChAT-positive neurons in the brainstem generally revealed immunopositivity for catecholaminergic markers. Local injections of neurotoxins in the substantia innominata blocked reticular facilitation, whereas local lesions of the septal nuclei and the nuclei of the diagonal band had no effect on MRF-induced facilitation. The blockage of the reticular facilitation of cortical evoked responses after unilateral lesions of the substantia innominata was bilateral, suggesting a cooperative interaction between basal forebrain structures of the two hemispheres. The anatomical and physiological data are discussed with respect to possible mechanisms of transient brainstem influences on cortical excitability. [ABSTRACT FROM AUTHOR]

The uniformly dense staining for cytochrome oxidase (CO) in layer IV of area 17 in adult monkeys has been described as developing from a bilaminar pattern in neonates being composed of a broad CO band in layer IVa and a narrow CO band attributed to the innermost zone of layer IV β. Here we present evidence, in the neonate marmoset monkey, for the narrow CO band to be localized in the outermost rim of layer V, designated as sublayer Va. [ABSTRACT FROM AUTHOR]

Focal blockade of restricted regions in layer V of area 18 was used to assess the contribution of this region to the responses to high-velocity stimuli of cells in retinotopically matched, layer V in area 17. In 40% of cases, blockade within area 18 revealed responses of area 17 cells to high-velocity stimuli to which they previously showed only poor responses. Stimulus specificity of the cells in area 17 was otherwise unaltered. All effects were reversible and repeatable. We suggest that a component of the output of layer V from area 18 normally suppresses the responses of retinotopically matched cells within area 17 to stimuli of high velocity, thereby enhancing the specificity of those cells to stimuli of low velocity [ABSTRACT FROM AUTHOR]

Radiofrequency lesions were made in the lateral geniculate nuclei of six squirrel monkeys. The resulting degenerating terminals and their postsynaptic structures in layer IV of area 17 were quantitatively categorized on photomontages covering large areas of neuropil. Two to five days after the lesion, numerous axon terminals were affected by a variety of degenerative changes, i.e., enlargement and distortion of synaptic vesicles, neurofilamentous hyperplasia, electron-lucent and electron-dense reactions. Based on the aggregation of electron-dense material beneath the postsynaptic membrane, the degenerating terminals were considered to be of the asymmetric type. Among the degenerating boutons were the largest axon endings that occur in layer IV. Three days postoperatively, degenerating boutons contributed an average of 16.2% to the total synapse population; five days postoperatively, the average had increased to 19.3 %. The percentage of degenerating boutons on individual montages, however, amounted to as much as 29%. This amount probably reflects more closely the actual contribution of the geniculocortical fiber system to layer IV of striate cortex. The postsynaptic structure most frequently contacted by degenerating axon endings was the dendritic spine, followed by dendrites of small diameter. To account for the diversity of degenerative changes in the same fiber system, we offer the tentative suggestion that heterogeneously degenerating axon terminals arise from a heterogeneous population of neurons in the lateral geniculate nucleus, i.e., from magnocellular versus parvocellular laminae. [ABSTRACT FROM AUTHOR]

Neurons and synapses were classified within layer IV of area 17 in monkeys ( Saimiri, Macaca mulatta, Macaca radiata). Three types of neurons of different sizes, distinct internal structure and synaptic environments occur. The first type is the largest neuron encountered in layer IV. It is rich in cytoplasmic organelles and receives a moderate number of Gray's type 1 and type 2 terminals onto soma and proximal dendrites. Type 1 endings are favored over type 2 terminals. This neuron resembles the large, smooth stellate cell of other investigators. The second type is a small neuron with fewer cytoplasmic organelles and few axosomatic endings of both types. It is similar to the small stellate cell described in other reports. The third type is the smallest cell in layer IV and contains only a sparse collection of organelles. It receives exclusively type 2 axon terminals onto soma and proximal dendrites and, thus, corresponds to LeVay's (1973) small spiny stellate cell. Neurons of this type are the most common in layer IV. Many of these small neurons contain an inclusion body. Gray's type 1 and type 2 axon terminals occur in different sizes. The most noteworthy terminals are type 1 endings of large size which establish many en passage contacts; they degenerate after destruction of the lateral geniculate nucleus. Large-sized type 2 terminals also make en passage contacts on:their tortuous course. The majority of both types of terminals are of small size. The ratio of type 1 to type 2 terminals is about 4∶1. The most common termination site of all classes of terminals is the dendritic spine, followed by small dendrites. The absence of presynaptic dendrites in layer IV suggests a fundamental difference between the wiring of visual cortex and subcortical visual structures and, thus, between the processing of visual information in cortex and lower centers. [ABSTRACT FROM AUTHOR]

Sensitivity to binocular positional mismatch was evaluated in cat striate cortical neurones, for paired, dichoptically presented, moving sine-wave gratings whose interocular positional phase relationship was varied. Spatial frequency, orientation and velocity were optimized for each neurone. Binocular responses for each spatial phase were compared with monocular stimulation of either eye. Binocular responses ranged from facilitation, through complete or partial summation, to partial or binocular occlusion. Counter to previous reports, all functional classes of neurones (simple neurones; special, intermediate and standard complex neurones) were represented in phase-specific and phase-insensitive groups. Most simple neurones, together with a small minority of standard complex neurones, exhibited near-total phase-sensitive modulation, the most significant new finding being that a minority of simple neurones were relatively insensitive to binocular mismatch. The majority of complex neurones, of all types, showed shallower modulation depths (typically around 30%), distributed in a continuum, with no indication of bimodality. It is concluded that the property of positional sensitivity is not attributable solely to simple neurones; and that positional insensitivity is not seen only in complex neurones. At least some neurones of all functional categories evince either kind of behaviour. These results are interpreted as a two-stage mechanism for convergence and matching of inputs from the two eyes. [ABSTRACT FROM AUTHOR]

Spatial-frequency dependence of directional tuning and directional bias was compared, for both eyes, in four previously established discrete classes of binocular feline striate cortical neurones. Two classes (respectively direction-selective or bidirectional at optimal spatial frequency) were directionality invariant at all spatial frequencies. In the remaining two classes, both directionbiased at optimal spatial frequency, directional bias either altered or reversed with change in spatial frequency. In all four classes, the directional tuning of a majority of neurones sharpened at high spatial frequency through either eye, although the bandpass characteristics were sometimes dissimilar for the two eyes. All neurones were of the same type through either eye. Amongst the two classes of direction-biased neurones, the strength of bias was commonly different through the two eyes. Where reversal of bias occurred, that reversal took place at different spatial frequencies for each eye. Thus, the direction and orientation preferences of cortical neurones are fixed at optimal spatial frequency, but their envelope of tuning to a gamut of spatial frequencies is not. These differences are potentially related to binocular coding of visual perspective, including dynamic object rotation in visual space. [ABSTRACT FROM AUTHOR]

The size and position of the guinea pig area 17 were determined by transneuronal labeling after intraocular injections of H-proline or WGA-HRP. Area 17 occupies a large region of the occipital cortex located between two shallow fissures, the fissura sagittalis lateralis and the lateral groove. Area 17 extends for about 6 mm rostral from the occipital pole of the hemisphere, and encroaches occipitally for more than 1 mm upon the ventromedial surface of the hemisphere; the lateral width is up to 4.5 mm. Single injections of WGA-HRP into area 17 produced eight patches of transported tracer which formed the same general pattern in the peristriate cortex, regardless of the position of the injection within the visual field representation of area 17. Two of these patches were found in anteromedial peristriate cortex; three patches were distributed anterolateral and lateral of area 17; and three patches were located in posterolateral peristriate cortex. For several reasons, each of these patches was interpreted as representing a single striate projection onto a separate peristriate area. Comparison of these results with published findings indicates that the parcellation of the peristriate cortex into a variety of different areas, the pattern formed by these areas around area 17, and their reciprocal connections with area 17 follow a common plan in all hitherto studied terrestrial Old World and New World rodents. Lucifer Yellow injections into striate cells projecting to one of the recipient areas (AM) indicated that the pyramidal cells of this set of striate neurons are characterized by a short apical dendrite, and that the basal dendrites of the layer V pyramidal cells branch more profusely than those of the layer III pyramids. [ABSTRACT FROM AUTHOR]

In eleven hemispheres of nine marmoset monkeys ( Callithrix jacchus), we have investigated the thalamo-cortical organization of the projections from the pulvinar to the striate and prestriate cortex. In each experiment, single or multiple injections of various retrograde fluorescent tracers were injected into adjacent regions or areas. In two experiments, horseradish peroxidase (HRP) was injected into the lateral geniculate nucleus (LGN) and the lateral pulvinar, respectively. The results show that the thalamo-cortical projection from LGN to striate cortex and from pulvinar to the prestriate cortex are similarly organized, but the geniculo-striate projection is more precise than the pulvinar-prestriate projection. The pulvinar-prestriate projection is topographically organized and preserves topological neighbourhood relations. Projection zones to the various visual areas are concentrically wrapped around each other. The projection zone to area 18 constitutes a central core region. It begins ventro-laterally in PuL where the pulvinar is in contact with the LGN. This contact zone we called the hilus region of the pulvinar. The area 18-projection zone stretches as a central cone into the posterior pulvinar through PuL and into PuM. It is surrounded by the projection zone to the posterior belt of area 19 and this in turn is surrounded by the projection zone to the anterior belt of area 19. The projection zones to area 19 are then surrounded medially and dorsally by zones projectiong to the temporal and parietal association cortex, respectively. The projection zone to area MT is located medio-ventrally in the posterior pulvinar (PuIP and surrounding nuclei) and coincides with a densely myelinated region. Area 17 also receives input from the pulvinar but probably predominantly in the region of the central visual field. The pulvinar zone projecting to area 17 is located ventrolaterally from the central core region projecting to area 18 and is contiguous laterally with the LGN. If the positions of the vertical and the horizontal meridian in the pulvinar correspond to those in the respective cortical projection zones, a second order visual field representation such as found in area 18, with the horizontal meridian split at an excentricity of about 7-10°, can also be recognized in the pulvinar. [ABSTRACT FROM AUTHOR]

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

The laminar distribution and postnatal development of profiles immunoreactive to antibodies directed against serotonin (5-hydroxytryptamine, 5-HT) have been investigated in the primary visual cortex (striate cortex, area 17) of cats. In the adult cat, profiles with serotonin-like immunoreactivity consist exclusively of fibers which exhibit laminar differences in density and predominant orientation. Immunoreactive fibers are dense in layers I-III, less dense in layer V, and sparse in layers IV and VI. In layers I and VI the trajectories of these fibers are mainly tangential to the pial surface; in layers II-V they are predominantly radial and more irregular. The vast majority of immunoreactive fibers consists of fine axons with frequent small varicosities. In addition, there are a few thick axons. In 2-week-old cats, immunoreactive fibers are sparsely distributed through layers II-V. By 4 weeks, fiber density has decreased still further in layer IV and increased in layers I-III. By 6 weeks, the laminar pattern resembles that of adult cats except that fiber density is still lower than in adults. At three months of age, the mature pattern is established. [ABSTRACT FROM AUTHOR]

Direction-selective or direction-biased striate cortical neurones were assessed for absence or incidence of suppression of firing, maximal at 90° or 180° ('null' suppression) to the optimal direction, in 327 neurones recorded from the striate cortex of cats anaesthetized with NO/O/halothane. Stimuli were light or dark bars moving over uniform or stationary textured back-grounds; or square-wave gratings of optimal spatial frequency and velocity. Five identified directionality groups were correlated with neuronal class and a range of other receptive field properties. Suppression maximal at 90° to optimum was common amongst direction-biased neurones, rare amongst direction-selective neurones. In the latter group, null suppression (maximal at 180° to optimum) was more prevalent than at 90°. Standard complex cells constituted the majority of complex neurones. They were more commonly direction-biased and less commonly showed suppression than special complex cells. The latter comprised the majority of direction-selective neurones with 180° suppression. Endstopping was seen more frequently in special complex cells, but for each functional class was similarly distributed between the different directionality groups. Based on the mean and mode of partially overlapping distributions, for all neuronal classes direction-selective neurones were more broadly tuned than direction-biased neurones. Special complex neurones were appreciably more broadly tuned than standard complex neurones; those with suppression at 180° were the most broadly tuned neurones in the cortex. Direction-biased neurones with suppression at 90° to optimum were more sharply tuned than those lacking such suppression. Direction-selective neurones had larger receptive fields than direction-biased neurones. In both groups receptive fields decreased in size in the sequence: intermediate complex > standard complex > special complex > simple. Resting discharge was highest amongst direction-selective neurones with 180° suppression, lower in those with 90° suppression or those lacking it, and lowest amongst direction-biased neurones. With the possible exception of the minority of neurones that were silent, low levels of resting discharge have not seriously prejudiced either neuronal categorization or comparisons of tuning selectivity. The pattern which emerges is that suppression maximal in directions orthogonal to the preferred direction/orientation is more commonly associated with sharp tuning and directionbias, whereas 'null' suppression, in the direction opposite that preferred, is associated with broad tuning, direction-selectivity, high resting discharge levels and strong texture sensitivity. [ABSTRACT FROM AUTHOR]

The numerical density of neurons and glial cells was estimated in visual area 18 of the adult human cerebral cortex and compared with that of area 17. Blocks of areas 17 and 18 came from the same brains and this allowed the comparison of 1) neuronal and glial numerical densities through the whole cortical depth with calculation of the neuron/glia ratio, 2) neuronal and glial numbers under one square millimeter of cortical surface, and 3) neuronal numerical densities in three groups of identified layers. The mean neuronal density is approximately 40000 neurons/mm in area 17 and 31500/m in area 18. The mean glial density is around 27000/mm in area 17 and 32000/mm in area 18. This gives a neuron/glia ratio of approximately 1.5 in area 17 and of 1.0 in area 18, but the total cellular density is similar in both areas. There are about 90000 neurons and 64000 glial cells under one square millimeter of cortical surface in area 17, and some 73000 neurons and 74000 glial cells in area 18. The higher neuronal density in area 17 is found through the whole depth of cortex and does not seem to be more pronounced in layer IVc of area 17 compared to layer IV in area 18 than in the groups of layers II-III and V-VI. [ABSTRACT FROM AUTHOR]

It is widely believed that cells in area 17 send axons specifically to neurons in other cortical areas whose receptive fields coincide with their own. We asked whether this was true in cats for area 17's projection to a large suprasylvian visual area, the Clare-Bishop area. Receptive fields were plotted at multiple sites in the Clare-Bishop area. Then, in area 17, anterograde tracer was injected at a retinotopically-characterized site, giving rise to patches of labeled terminals in the Clare-Bishop area. Receptive field centers recorded within these patches were located close to the visual field location at the injection site in area 17. Receptive fields recorded outside of labeled patches, on the other hand, were never in register with that plotted in area 17. However, due to their large size, even fields located outside of labeled patches often encompassed the visual field point injected in area 17. In other experiments, receptive fields for both neurons and presumed cortical afferents were recorded at the same site in the Clare-Bishop area. The centers of such pairs of receptive fields were on average less than 1° apart. Finally, the gaps between widely separated patches of label were investigated. Both physiological and anatomical evidence indicated that a different part of the visual field was represented in gaps than in the adjacent patches. [ABSTRACT FROM AUTHOR]

The postnatal development of the striate cortical projection onto the extrageniculate visual thalamus was examined by using the orthograde and retrograde HRP methods. At birth, the projection is present, and fibres of the projection terminate in the lateral part of the lateral posterior nucleus. A rough topographical arrangement of the projection is already established. Neurones that give rise to the projection are located exclusively in layer V from birth onward. However, the neurones are much more densely packed at birth than in adult cats. The packing density of the neurones decreases rapidly during the second postnatal week, and afterwards continues to become gradually lower. In the meantime, the cross sectional area of the neurones increases sharply during the second postnatal week. These findings suggest that striate cortical neurones projecting onto the lateral posterior nucleus rapidly complete the final stages of their maturation shortly after the normal opening of the eyelids, and during this time some of these neurones undergo axonal elimination or neuronal death, or both. [ABSTRACT FROM AUTHOR]

Recordings from single units in kitten primary visual cortex show that a reversible blockade of the discharge activities of cortical neurons and geniculocortical afferent terminals by intracortical infusion of the sodium channel blocker tetrodotoxin (TTX) completely prevented the ocular dominance shift that would normally be seen after monocular deprivation. The blockade of cortical plasticity, like the blockade of discharge activity, was reversible, and plasticity was restored following recovery from the effects of TTX. These results extend previous work suggesting involvement of electrical activity at the level of the cortex in the phenomenon of cortical plasticity by demonstrating an absolute requirement for discharge activities in the primary visual cortex. [ABSTRACT FROM AUTHOR]

Single units were recorded in area 17 of anesthetized and paralyzed cats. The discharges of cortical units, either spontaneous or driven by two dimensional visual drifting noise, were analyzed during unilateral tactile stimulation (air puffs or taps) of the skin around the eye. This stimulation evokes a blink response in the normal non-curarized animal. The activity in the branch of the facial nerve innervating the orbicularis oculi muscles responsible for the blink was also recorded on the stimulated side. Following the mechanical stimulation, the discharges of both simple and complex visual cells were strongly inhibited with a latency of 70-80 ms. Inhibition was sometimes preceded by a brief increase in firing rate. This typical response was present only when the cutaneous stimulus was effective in triggering a discharge in the motor nerve which drives the orbicularis oculi muscles. Moreover, when a visual response was evoked by a temporary masking of the visual stimulus, this response was suppressed by the association with a tap stimulation. [ABSTRACT FROM AUTHOR]

The developmental properties of 573 neurones have been investigated in the primary visual cortex of eight binocularly intact and twelve unilaterally enucleated kittens. It is shown that removal of one eye at birth alters the development of orientation selectivity observed in the presence or absence of visual experience. In 6-week-old deprived kittens, there remain significantly more orientation selective cells in enucleated than in binocularly deprived kittens. These deprivation-resistant cells respond preferentially to horizontal or vertical orientations and are recorded mainly in the cortex contralateral to the remaining eye. In six-week-old kittens with visual experience, the process of tuning maturation appears to be unaffected by unilateral enucleation at birth. However, a larger over-representation of horizontal and vertical orientation preferences is observed in uniocular kittens than in binocularly intact kittens, suggesting that the development of oblique orientation preference depends upon the presence of binocular afferents in the visual pathway. [ABSTRACT FROM AUTHOR]

Small electrophoretic injections of horseradish peroxidase (HRP) were made with recording microelectrodes in different layers of area 17 of the cat. The sizes and locations of labeled relay cells in the lateral geniculate nucleus (LGNd) were then studied. Layer I was found to receive inputs mainly from small LGNd cells (13-21 μm). These cells were located in the parvocellular C laminae and the MIN. Layer II and the upper portion of layer III were not found to receive LGNd afferents. Lower layer III was found to receive inputs mainly from small to medium-sized relay cells. These cells were located in the A laminae and the parvocellular C laminae. Layer IVab and the layer III-IV border region were found to receive afferents from predominantly large LGNd neurons (> 20 μm). These cells were located in laminae A, A; C, and the MIN. Layer IVc was found to receive inputs from mostly small to medium-sized cells (15-23 μm). Nearly all of these cells were in the A laminae although some small cells in laminae C were observed following layer IVc injections which involved the layer IV-V border region. Layer V, away from the layer IV-V border region, was not found to receive LGNd afferents. Finally, layer VI was found to receive inputs from both large and small relay cells. These cells were all located in the A laminae. In the light of previous findings, these results suggest that the terminals of W, X, and Y cell axons are segregated within the cat's striate cortex: W cells projecting to layer 1, the deeper portion of layer III and the layer IV-V border region; X cells projecting heavily to layer IVc with a minor projection to layer VI and possibly also to lower layer III; Y cells projecting heavily to layer IVab and the layer III-IV border region with a smaller projection to layer VI. [ABSTRACT FROM AUTHOR]

Neuronal responses to moving visual stimuli were recorded in the lateral geniculate nucleus (LGN) and area 17 of cats. Response duration (D), number of spikes (N), and mean frequency (F) were estimated from the response histograms and analysed for their dependence on stimulus velocity. In the LGN, for about 2/3 of cells these response parameters changed monotonically with velocity up to about 100 °/s. In 1/3 of the cells, the response frequency was tuned to velocity. The speed at which individual cells reached a peak or plateau firing rate was correlated with their receptive field size. In area 17, most neurones were tuned to velocity. Nine out of 59 cells were insensitive to stimulus speed in that they responded equally well at stimulus velocities up to about 10 °/s. The results suggest that at higher levels in the nervous system information about velocity is represented in discrete groups of neurones. It is pointed out that different response parameters may be relevant for different perceptual phenomena associated with movement. The significance of integrational properties and lateral inhibition of nerve cells for the development of complex response properties is discussed. [ABSTRACT FROM AUTHOR]

The response of single cells in the striate cortex of cat to a moving light bar of variable orientation was measured by a method providing data on the mean response as well as the standard deviation (SD) at the different stimulus orientations. At the optimal stimulus orientation the SD was about 1/3 of the mean response. Marked differences in this respect were found between simple and complex cells, i.e., the SD for the simple cells was about 1/2 of the mean response and about 1/4 for the complex cells. The variation coefficient (Vc = SD/mean) was minimal at the optimal orientation and increased relatively in the same manner for simple and complex cells as the stimulus orientation was varied away from optimal orientation. The Vc varied with the mean response at optimal orientation in a nonlinear manner. A function is proposed which fits this relationship and which is equally applicable for both simple and complex cells. The mean orientation discrimination (MOD) was defined as that change in orientation angle away from the optimal which produced a response statistically different - on the 1 % level - from the response to the optimal orientation. There were differences in MOD between the two sides of the orientation tuning curve: the mean of the smaller of the two values was 13.5 deg and of the larger 19.7 deg. No significant difference in MOD was found between simple and complex cells despite the fact that the halfwidth of the tuning curves for the two cell types was 19.5 deg and 31.6 deg, respectively. The preciseness in localization of the most sensitive part within the receptive field of single cells was calculated from the variability in time of occurrence of the smallest interspike interval. The degree of preciseness was found to be of the order of 1/4 of the receptive field diameter in both simple and complex cells. When nonoptimal stimulus orientations were presented, the preciseness significantly decreased in complex cells whereas it remained unchanged in simple cells. It is suggested that the same type of intracortical wiring produces orientation selectivity in simple and complex cells, and that the differences in tuning width are mainly due to a larger extension of inhibitory fields in the simple cells. Considering the cortical visual cells as elementary units in a network built for orientation detection and discrimination, the tuning width seems of minor importance for that function. [ABSTRACT FROM AUTHOR]

Single units were recorded extracellularly from area 17 and lateral suprasylvian area (LSSA) in curarized cats. Visual stimuli, usually a 10 ° black spot, were introduced abruptly in the visual field remote from the discharge area of a neuron's receptive field and moved at a speed of about 30 °/sec. The effect of these remote stimuli (S) on the response to a restricted visual stimulus (S) crossing the discharge area was studied. It was found that most units in area 17 were not affected by the presentation of remote stimuli, the remainder being either slightly facilitated or slightly inhibited. In contrast the LSSA neurons were usually inhibited by the presentation of S: this effect was strong, was present in all classes of LSSA neurons and was independent of the relative directions of movement of S and S. On the basis of these data and those previously obtained from the superior colliculus it is concluded that the way the extrageniculate centres respond to a stimulus abruptly introduced in the visual field is substantially different from that of the striate cortex. Only in the extrageniculate centres a new stimulus, besides exciting the neurons which correspond to the position of the stimulus in the field, concomitantly decreases the responses of neurons located in positions of the visual field remote from that stimulus. Possible behavioral implications of the findings are discussed. [ABSTRACT FROM AUTHOR]

Cells in cat's area 17 respond optimally if elongated contrasts are presented at a certain angle of orientation with respect to the retina, or to the visual field, respectively (Hubel and Wiesel, 1962). The preferred orientation and the range of orientation sensitivity of cells in close proximity to one another have been determined in order to investigate the spatial arrangement of the orientation domain in area 17. [ABSTRACT FROM AUTHOR]

In principle, the development of sensory receptive fields in cortex could arise from experience-independent mechanisms that have been acquired through evolution, or through an online analysis of the sensory experience of the individual animal. Here we review recent experiments that suggest that the development of direction selectivity in carnivore visual cortex requires experience, but also suggest that the experience of an individual animal cannot greatly influence the parameters of the direction tuning that emerges, including direction angle preference and speed tuning. The direction angle preference that a neuron will acquire can be predicted from small initial biases that are present in the naïve cortex prior to the onset of visual experience. Further, experience with stimuli that move at slow or fast speeds does not alter the speed tuning properties of direction-selective neurons, suggesting that speed tuning preferences are built in. Finally, unpatterned optogenetic activation of the cortex over a period of a few hours is sufficient to produce the rapid emergence of direction selectivity in the naïve ferret cortex, suggesting that information about the direction angle preference that cells will acquire must already be present in the cortical circuit prior to experience. These results are consistent with the idea that experience has a permissive influence on the development of direction selectivity. [ABSTRACT FROM AUTHOR]