Visual responses of neurons in the avian nucleus isthmi

Electrophysiological responses of neurons to visual and auditory stimulation are extracellularly recorded from the pigeon isthmic area. Cobalt sulfide markings show that only visual units are localized within the nucleus isthmi pars parvocellularis (Ipc) and pars magnocellularis (Imc), while visual-auditory bimodal units are localized outside. Visual units respond to black or white targets moving through their receptive fields (RFs). The RF centers are mainly distributed in the contralaterally lower visual field. The rostral Ipc and Imc receive information from the nasal visual field, and the caudal part of the Ipc and Imc corresponds to the temporal field. Therefore, both Ipc and Imc are visual centers instead of auditory centers as described before.     

Magnocellular and parvocellular divisions of pigeon nucleus isthmi differentially modulate visual responses in the tectum

The electrophysiological responses of 162 tectal cells to computer-generated visual stimuli were extracellularly recorded from 24 homing pigeons before and after injecting either lidocaine or N-methyl-d-aspartate (NMDA) into the nucleus isthmi pars magnocellularis (Imc) or the nucleus isthmi pars parvocellularis (Ipc). Micro-injections of lidocaine into Imc resulted in a significant reduction of firing rate in 80% of tectal cells, whose excitatory receptive fields (ERFs) were localized within the ERF of the Imc cell where the lidocaine was injected. In contrast, when lidocaine was injected into Ipc under identical circumstances it had no effect on the visually driven activity of 68% of tectal cells. However, when the excitatory amino acid NMDA was injected into Ipc it produced a significant reduction in the visually driven firing of 75% of tectal neurons when their ERFs were within the isthmic ERF, while similar application of NMDA into Imc had no effect on the visually driven response of 94% of tectal neurons. When the ERFs of tectal cells were localized outside the ERF of the isthmic cell where the chemical was injected, Imc-injected lidocaine had no effect in 9 out of 10 tectal cells, whereas Ipc-injected NMDA increased firing in 7 out of 17 tectal cells. Therefore, it is suggested that the Imc-tectal fibers participate in a positive feedback pathway and the Ipc-tectal fibers are involved in a negative feedback pathway.     

Visual experience as a determinant of the response characteristics of cortical receptive fields in cats

Summary Cats reared with their visual world restricted to vertical lines for one eye and horizontal lines for the other had, in their visual cortices, units with elongated receptive fields that were either vertically or horizontally oriented. These receptive fields could be mapped only using that eye which had seen lines of the same orientation during development. Other units had diffuse, unresponsive receptive fields (Hirsch and Spinelli, 1970). Six cats, from the group above, were revived and allowed normal binocular viewing in an attempt to determine the possibility and extent of adding other types of receptive fields by giving other experiences to their visual systems. After exposure to a normal environment for up to 19 months it was found that indeed there had been a massive increase in the percentage of those classes of receptive fields that were either absent or weak at the end of the selective visual experience. Significantly, these receptive fields, acquired during binocular viewing, were very often binocular.The results, however, show that units whose response characteristics mimic the stimuli viewed during development were almost completely unaffected by normal binocular visual experience, i. e., they were monocularly activated and had the orientation appropriate for the stimuli viewed by the eye from which they could be mapped. Most impressive are a few units whose receptive field shape is almost a carbon copy of the pattern viewed during development. The data provide evidence that visual experience has a direct continuing and lasting effect on the functional connectivity of cells in the visual cortex.     

Diverse receptive fields in the lateral geniculate nucleus during thalamocortical development

Most models of thalamocortical development in the visual system assume a homogeneous population of thalamic inputs to the cortex, each with concentric on- or off-center receptive fields. To test this, we made high-resolution spatial maps of receptive fields in the developing ferret lateral geniculate nucleus (LGN). Developing receptive fields (RFs), had a variety of shapes: some concentric, others elongated (like adult cortical receptive fields) and some with 'hot spots' of sensitivity. These receptive fields seemed to arise from convergence of multiple retinal afferents onto LGN neurons. We present a Hebbian model whereby imprecise retinogeniculate connections help refine geniculocortical connections, sharpening both thalamocortical topography and perhaps orientation selectivity.     

Neurons with large bilateral receptive fields in monkey prelunate gyrus

In single-cell recordings from the dorsocaudal part of the prelunate gyrus of an alert monkey (Macaca fascicularis) we found neurons with unexpectedly large receptive fields (RFs) that spread bilaterally into the contra- and ipsilateral visual fields. These neurons (n=82) appeared to be clustered in the periphery of V4. They were surrounded by neurons with relatively small (3-10 degrees) and unilateral RFs in the contralateral field with properties similar to those previously described for neurons in area V4. Bilateral RFs extended over large parts of the lower visual field but always spared the fovea. Receptive fields typically revealed two foci of maximal responsiveness that were arranged symmetrically in the ipsi- and contralateral fields. Twenty-six cells did not respond to stimuli along the vertical meridian; these neurons had two distinct RFs. The preference for stimulus orientation, color, or motion was similar in all parts of these large RFs.     

Distant cortical locations of the upper and lower quadrants of the visual field represented by neurons with elongated and radially oriented receptive fields

In our previous study of the cytoarchitectonic field 7 of cat cortex we had described neurons with extremely elongated receptive fields (RFs). The long axes of these RFs were oriented radially, towards the centre of the retina. These neurons represented only the lower contralateral part of visual field. They were surrounded from all sides by neurons with clearly different RF properties. We proposed that neurons with a similar radial organization and with RFs in the upper visual field also exist in the cortex but are localized in the area that was distant from the representation of the corresponding lower visual field. We expected to find these neurons in front of the representation of the upper visual field in areas V1, V2 and V3 (fields 17, 18 and 19), behind the central representation in area 21a. This cortical region was studied in five behaving cats. In all animals, neurons with radial RFs in the upper visual field were found in the expected location. As in the lower visual field, their RFs always spared the central visual field. Other RF properties of these neurons were also very similar to those found previously in the lower visual field. It became obvious that neurons with radial RFs are included into the fourth extrastriate crescent with complete contralateral representation. However, in the fourth crescent, RF properties in the central visual field differed significantly from those on the periphery. As a result, neurons with similar radial RFs in the upper and lower visual fields were located in the distant cortical regions, and were separated by the representation of the central visual field presented by the non-radial neurons of the cytoarchitectonic area 21a.     

Unusually large receptive fields in cats with restricted visual experience

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

Cat area 17. III. Response properties and orientation anisotropies of corticotectal cells

The receptive field properties of antidromically identified corticotectal (CT) cells in area 17 were explored in the paralyzed, anesthetized cat. To compare these with another population of infragranular cells, we also examined the receptive field properties of cells in layer 6. Sixty percent of our sample of CT cells showed increased response to increased stimulus length (length summation) and were classified as standard complex cells. The other 40% showed little or no length summation, were generally end stopped, and were classified as special complex cells. Standard and special complex CT cells have complementary orientation anisotropies: the distribution of orientation preferences of standard complex cells is biased toward obliquely oriented stimuli, whereas special complex cells are biased toward horizontally and vertically oriented stimuli. The receptive fields of the cells in our sample were primarily along the horizontal meridian so we cannot determine if these anisotropies are defined relative to the vertical meridian or relative to the meridian passing through the receptive field. The effects of these anisotropies in preferred orientation are minimized by the broad orientation tuning of CT cells. There was no simple relationship between the direction bias of CT cells and the reported direction bias of tectal cells. In contrast to the heterogeneity of corticotectal cells, layer 6 cells uniformly showed strong length summation, tight orientation tuning, and little spontaneous activity.     

The effects of aging on the strength of surround suppression of receptive field of V1 cells in monkeys

The surround suppression of the receptive field is important for basic visual information processing, such as orientation specificity. To date, the effects of aging on the strength of surround suppression are not clear. To address this issue, we carried out extracellular single-unit studies of the receptive field properties of cells in the primary visual cortex (area V1) in young and old rhesus (Macaca mulatta) monkeys. When presented with the oriented central stimulus, we found that cells in old animals showed reduced orientation and direction selectivity compared with those in young animals. When presented with the oriented central stimulus together with the optimal surround stimulus, more selective cells {orientation bias (OB) ≥0.1; a bias of 0.1 is significant at the P<0.005 level} in animals of both ages showed reduced orientation selectivity compared with the experiment that presented only the oriented central stimulus. When presented with the optimal central stimulus together with the oriented surround stimulus, cells in old animals showed reduced orientation and direction selectivity compared with young animals. Moreover, broadly tuned cells (OB<0.1) in old animals exhibited significantly reduced suppression indices that quantified the strength of the surround suppression of the receptive field, when compared with those in young animals. These results suggest that aging may seriously affect the surround suppression of the receptive field of V1 cells. Thus, the decreased strength of surround suppression of the receptive field may be one possible reason for the decreased stimulus selectivity of V1 cells previously found in the senescent brain. This work will contribute to an understanding of the physiological mechanisms mediating surround suppression of the receptive field.     

The primary visual cortex in the mouse: Receptive field properties and functional organization

Summary Receptive field (RF) characteristics of cells in primary visual cortex of the mouse (C57B16 strain) were studied by single unit recording. We have studied the functional organization of area 17 along both the radial and tangential dimensions of the cortex. Eighty seven percent of the visual neurons could be classified according to their responses to oriented stimuli and to moving stimuli. Cells which preferred a flashed or moving bar of a particular orientation and responded less well to bars of other orientations or to spots, were classified as orientation selective (simple RF 23%, complex RF 18%). The majority of them were, moreover, unidirectional (24%). All orientations were roughly equally represented. Cells with oriented RFs were recorded mostly in the upper part of cortical layers II–III, where they appeared to be clustered according to their preferred orientation. Neurons that responded equally well to spots and bars of all orientations (46%) were classified as non-oriented; among these neurons there were several subcategories. Cells which responded equally well to spots and bars but preferred stimuli moving along one or both directions of a particular axis were classified as non oriented asymmetric cells (unidirectional 14%, bidirectional 4%). They were recorded mainly in supra- and infra-granular layers. Cells unaffected by stimulus shape and orientation which responded equally well to all directions of movement were classified as symmetric units. They had receptive field classified as ON (11%), OFF (1%), ON/ OFF (11%), or were unresponsive to stationary stimuli (5%). These cells were mostly found in layer IV, in which they constituted the majority of recorded cells. There was no apparent correlation between the functional type and size of RFs. However, the greatest proportion of small RFs was found in layer IV. In the binocular segment of the mouse striate cortex, the influence of the contralateral eye predominated. Ninety five percent of cells in this segment were driven through the contralateral eye. However, 70% of cells were binocularly activated, showing that considerable binocular integration occured in this cortical segment. Ocular dominance varied less along the radial than along the tangential dimension of the cortex.     

Cat retinal ganglion cells: size and shape of receptive field centres

1. Receptive field centres of 144 sustained and transient retinal ganglion cells were mapped in cats under light pentobarbitone anaesthesia.2. Sustained on-centre, sustained off-centre, transient on-centre and transient off-centre cells had different mean sizes of receptive field centre, with some overlap between their distributions.3. For each class of cell, central fields had the smallest field-centres; progressively larger field-centres were encountered more peripherally.4. All classes of ganglion cells tended to have slightly elliptical receptive field centres. Major axes of over half of all receptive fields were oriented within 20 degrees of horizontal. These trends were independent of pupil dimensions, or of receptive field eccentricity or position in the visual field. The results almost certainly reflect asymmetry in retinal wiring.5. Two cells of thirty-nine tested were sensitive to axis of motion; in both cases the preferred and major axis were horizontal. A further cell was orientation specific.     

Visual perception in cats after environmental surgery

Summary This report describes a series of experiments testing the behavioral significance of orientation sensitive cells in the cat's visual cortex. Cats used for this purpose were raised with one eye (VE) viewing a field of vertical lines and the other eye (HE) viewing a field of horizontal lines. This experience simplified their cortical physiology, and is thus called environmental surgery. Cells with elongated fields were found to be activated monocularly: those driven by the VE had vertically oriented receptive fields, while those activated by the HE had horizontally oriented fields. These animals were tested, using one eye at a time, on a series of visual discriminations: a) flux discrimination, b) tests for selective response to vertical and horizontal lines, c) discrimination between two lines differing only in orientation, d) discrimination between mirror image stimuli, and e) evaluation of inter-ocular transfer on these discriminations. The threshold tests for orientation discrimination gave the clearest results: there were small but consistent differences in performance between the VE and HE which depended on the orientation of the lines being discriminated. There was also evidence that the animals responded to different parts of a stimulus with the VE and HE. As a group, the experimental cats showed less interocular transfer of visual discriminations than the normal controls. However, surprisingly, there was also evidence for much functional equivalence between the two eyes. Possible explanations for this are considered and it is suggested that an animal's ability to make pattern discriminations is not rigidly determined by the shape and orientation of its receptive fields.     

Pulvinar nuclei of the behaving rhesus monkey: visual responses and their modulation

We have examined the properties of neurons in three subdivisions of the pulvinar of alert, trained rhesus monkeys 1) an inferior, retinotopically mapped area (PI), 2) a lateral, retinotopically organized region (PL), and 3) a dorsomedial visual portion of the lateral pulvinar (Pdm), which has a crude retinotopic organization. We tested the neurons for visual responses to stationary and moving stimuli and for changes in these responses produced by behavioral manipulations. All areas contain cells sensitive to stimulus orientation as well as neurons selective for the direction of stimulus movement; however, the majority of cells in all three regions are either broadly tuned or nonselective for these attributes. Nearly all cells respond to stimulus onset, a significant number also give a response to stimulus termination, and rarely a cell gives only off responses. Nearly all cells increase their discharge rate to visual stimuli. Receptive fields in the two retinotopically mapped regions, PI and PL, have well-defined borders. The sizes of these receptive fields show a positive correlation with the eccentricity of the receptive fields. The receptive fields in the remaining region, Pdm, are frequently very large, but with these large fields excluded, show a similar correlation with eccentricity. All pulvinar cells tested (n = 20) were mapped in retinal coordinates; the receptive fields are positioned in relation to the retina. We found no cells with gaze-gated characteristics (2), nor cells mapped in a spatial coordinate system. The response latencies in PI and PL are shorter and less variable than the latencies in Pdm. Active use of a stimulus can produce an enhancement or attenuation of the visual response. Eye-movement modulation was found in all three subdivisions in about equal frequencies. Attentional modulation was common in Pdm and was rare in PI and PL. The modulation is spatially selective in Pdm and nonselective in PI for a small number of tested cells. These data demonstrate functional differences between Pdm and the other two areas and suggest that Pdm plays a role in selective visual attention, whereas PI and PL probably contribute to other aspects of visual perception.     

The spatial extent of excitatory and inhibitory zones in the receptive field of superficial layer hypercomplex cells

1. An investigation has been made of the extent of inhibitory and excitatory components in the receptive field of superficial layer hypercomplex cells in the cat's striate cortex and the relation of the components to the length preference exhibited by these cells.2. Maximal responses were produced by an optimal length stimulus moving through a restricted region of the receptive field. The length of this receptive field region was less than the total length of the excitatory zone as mapped with a very short slit. Slits of similar length to the excitatory zone produced a smaller response than an optimal length slit.3. An increase of slit length so that it passed over receptive field regions either side of the excitatory zone resulted in an elimination of the response. When background discharge levels were increased by the iontophoretic application of D, L-homocysteic acid slits of this length were observed to produce a suppression of the resting discharge as they passed over the receptive field. They did not modify the resting discharge level when it was induced by the iontophoretic application of the GABA antagonist bicuculline. This data is taken to indicate that long slits activate a powerful post-synaptic inhibitory input to the cell.4. Maximal inhibitory effects were only observed if the testing slit passed over the receptive field centre. That is slits with a gap positioned midway along their length so as to exclude the optimal excitatory response region surprisingly tended to produce excitatory effects rather than the expected inhibitory effects. It appears that simultaneous stimulation of the receptive field centre is a precondition for the inhibitory effect of stimulation of regions either side of the excitatory zone to be activated.5. It is suggested that the interneurones mediating the inhibitory input to the superficial layer hypercomplex cells are driven both by cells in adjacent hypercolumns with receptive fields spatially displaced to either side of the excitatory zone and by cells in the same column, optimal inhibitory effects only being achieved when both sets of input to the interneurone are activated.     

Visual receptive-field properties of cells in area 18 of cat's cerebral cortex before and after acute lesions in area 17.

1. Receptive-field properties of single neurons in cat's cortical area 18 were studied before and after partial bilateral lesions of area 17. 2. The majority of cells recorded from animals with intact visual cortex exhibited orientation selectivity, directional selectivity, and could be independently activated through either eye. All cells responded well to moving targets and nearly all of them exhibited broadly tuned preferences with respect to speed of the target. Over 45% of cells responded optimally or exclusively at very fast (above 50 degrees/s) speeds. 3. The majority of neurons recorded from animals with intact visual cortex responded weakly but clearly to appropriately oriented localized stationary stimuli flashed on and off. About one-third of the cells responded with mixed on-off discharges from all over their receptive field. In the receptive fields of 10% of cells, separate on- and off-discharge regions could be revealed. In the receptive fields of the remaining cells, only on- or only off-discharge regions could be revealed. 4. The majority of neurons recorded after ablation of area 17 were orientation selective; 50% of the cells were also direction selective. All neurons responded well to moving targets; about 65% of them responded optimally or exclusively at very fast target speeds. 5. Destruction of the dorsolateral part of contralaterial area 17 and most of contralateral area 18 caused significant reduction in proportion of cells in area 18 which could be activated through either eye. 6. The majority of neurons recorded after ablation responded to appropriately oriented localized stationary stimuli flashed on and off. Cells with mixed on-off discharge regions all over the receptive field with separate on- and off-discharge regions and with only on- or only off-discharge regions were found. 7. It is concluded that the processing of afferent visual information in area 18 is, to a great extent, independent of the information carried to this area by associational fibers from cells of area 17.     

Spatio-temporal plasticity of cortical receptive fields in response to repetitive visual stimulation in the adult cat

Many psychophysical experiments on perceptual learning in humans show increases of performance that are most probably based on functions of early visual cortical areas. Long-term plasticity of the primary visual cortex has so far been shown in vivo with the use of visual stimuli paired with electrical or pharmacological stimulation at the cellular level. Here, we report that plasticity in the adult visual cortex can be achieved by repetitive visual stimulation. First, spatial receptive field profiles of single units (n=38) in area 17 or 18 of the anesthetized cat were determined with optimally oriented flashing light bars. Then a conditioning protocol was applied to induce associative synaptic plasticity. The receptive field center and an unresponsive region just outside the excitatory receptive field were synchronously stimulated ('costimulation', repetition rate 1 Hz; for 10-75 min). After costimulation the receptive field and its adjacent regions were mapped again. We observed specific increases of the receptive field size, changes of the receptive field subfield structure as well as shifts in response latency.In 37% of the cells the receptive field size increased specifically towards the stimulated side but not towards the non-stimulated opposite side of the receptive field. In addition, changes in the relative strength and size of the on and off subfield regions were observed. These specific alterations were dependent on the level of neuronal activity during costimulation. During recovery, the new responses dropped down to 120% of the preconditioning value on average within 103 min; however, the decay times significantly depended on the response magnitude after costimulation. In the temporal domain, the latency of new responses appeared to be strongly influenced by the latency of the response during costimulation.Twenty-nine percent of the units displayed no receptive field enlargement, most likely because the activity during costimulation was significantly lower than in the cases with enlarged receptive fields. An unspecific receptive field enlargement towards both the stimulated and non-stimulated side was observed in 34% of the tested cells. In contrast to the cells with specifically enlarged receptive fields, the unspecific increase of receptive field size was always accompanied by a strong increase of the general activity level.We conclude that the receptive field changes presumably took place by strengthening of synaptic inputs at the recorded cells in a Hebbian way as previously shown in the visual cortex in vitro and in vivo. The observed receptive field changes may be related to preattentive perceptual learning and could represent a basis of the 'filling in' of cortical scotomas obtained with specific training procedures in human patients suffering from visual cortex lesions.     

蜥蜴中脑神经通路和起源细胞的形态

本文采用 HRP 法研究了蛤蚧(Gekko gekko)和鳄蜥(Shinisaurus crocodilurus)视顶盖、中脑深核(NPM)与峡核之间的通路和起源细胞的形态。结果指出:1.顶盖与峡核大细胞部(Imc)呈相互区域对应投射;2.同侧顶盖—Imc 投射细胞主要位于第7层,系有径向树突的梨形细胞;同侧 Imc—顶盖投射细胞为小树突域的梨形或多角形细胞;3.顶盖注射标记的 NPM细胞呈纺锤形,染色浅;峡核注射标记的 NPM 细胞,其粗树突往往伸向顶盖;4.NPM 注射标记顶盖细胞和峡核细胞,前者主要位于顶盖第7层,后者散布在峡核大细胞部(Imc)和峡核小细胞部(Ipc)内。     

GABA-inactivation attenuates colinear facilitation in cat primary visual cortex

Neurons in primary visual cortex (V1) respond preferentially to stimuli of a particular orientation falling within a circumscribed region of visual space known as their receptive field (RF). However, the response to an optimally oriented stimulus presented within the RF can be enhanced by the simultaneous presentation of co-oriented, co-linearly aligned flank stimuli falling outside the RF which, when presented alone, fail to activate the cell. This type of contextual effect, termed colinear facilitation, presumably forms the physiological substrate for the integration of the line elements of a contour and the perceptual saliency of a contour in a complex environment. Here we show that colinear facilitation in single cells of cat area V1 can be substantially reduced or abolished by focal inactivation of laterally remote cells in the same area which respond strongly to the co-oriented, colinear flank stimulus inducing the facilitatory effect. The results provide evidence that horizontal intrinsic connections between cells with co-oriented and co-linearly aligned RFs make a major contribution to colinear facilitation in V1. They imply that the neuronal circuitry underlying contour integration and saliency is already present at the earliest stage of visual cortical information processing.     

Receptive fields and functional architecture of monkey striate cortex

1. The striate cortex was studied in lightly anaesthetized macaque and spider monkeys by recording extracellularly from single units and stimulating the retinas with spots or patterns of light. Most cells can be categorized as simple, complex, or hypercomplex, with response properties very similar to those previously described in the cat. On the average, however, receptive fields are smaller, and there is a greater sensitivity to changes in stimulus orientation. A small proportion of the cells are colour coded.

2. Evidence is presented for at least two independent systems of columns extending vertically from surface to white matter. Columns of the first type contain cells with common receptive-field orientations. They are similar to the orientation columns described in the cat, but are probably smaller in cross-sectional area. In the second system cells are aggregated into columns according to eye preference. The ocular dominance columns are larger than the orientation columns, and the two sets of boundaries seem to be independent.

3. There is a tendency for cells to be grouped according to symmetry of responses to movement; in some regions the cells respond equally well to the two opposite directions of movement of a line, but other regions contain a mixture of cells favouring one direction and cells favouring the other.

4. A horizontal organization corresponding to the cortical layering can also be discerned. The upper layers (II and the upper two-thirds of III) contain complex and hypercomplex cells, but simple cells are virtually absent. The cells are mostly binocularly driven. Simple cells are found deep in layer III, and in IV A and IV B. In layer IV B they form a large proportion of the population, whereas complex cells are rare. In layers IV A and IV B one finds units lacking orientation specificity; it is not clear whether these are cell bodies or axons of geniculate cells. In layer IV most cells are driven by one eye only; this layer consists of a mosaic with cells of some regions responding to one eye only, those of other regions responding to the other eye. Layers V and VI contain mostly complex and hypercomplex cells, binocularly driven.

5. The cortex is seen as a system organized vertically and horizontally in entirely different ways. In the vertical system (in which cells lying along a vertical line in the cortex have common features) stimulus dimensions such as retinal position, line orientation, ocular dominance, and perhaps directionality of movement, are mapped in sets of superimposed but independent mosaics. The horizontal system segregates cells in layers by hierarchical orders, the lowest orders (simple cells monocularly driven) located in and near layer IV, the higher orders in the upper and lower layers.

     

Polysensory properties of neurons in the anterior bank of the caudal superior temporal sulcus of the macaque monkey

1. We examined the sensory properties of cells in the anterior bank of the caudal part of the superior temporal sulcus (caudal STS) in anesthetized, paralyzed monkeys to visual, auditory, and somesthetic stimuli. 2. In the anterior bank of the caudal STS, there were three regions distinguishable from each other and also from the middle temporal area (MT) in the floor of the STS and area Tpt in the superior temporal gyrus. The three regions were located approximately in the respective inner, middle, and outer thirds of the anterior bank of the caudal STS. These three regions are referred to, from the inner to the outer, as the medial superior temporal region (MST), the mostly unresponsive region, and the caudal STS polysensory region (cSTP), respectively. 3. The extent of MST and its response properties agreed with previous studies. Cells in MST responded exclusively to visual stimuli, had large visual receptive fields (RFs), and nearly all (91%) showed directional selectivity. 4. In the mostly unresponsive region, three quarters of cells were unresponsive to any stimulus used in this study. A quarter of the cells responded to only visual stimuli and most did not show directional selectivity for moving stimuli. Several directionally selective cells responded to movements of three-dimensional objects, but not of projected stimuli. 5. The response properties of cells in the superficial cortex of the caudal superior temporal gyrus, a part of area Tpt, external to cSTP were different from those of cells in the three regions in the anterior bank of the STS. Cells in Tpt were exclusively auditory, and had much larger auditory RFs (mean = 271 degrees) than those of acoustically-driven cSTP cells (mean = 138 degrees). 6. The cSTP contained unimodal visual, auditory, and somesthetic cells as well as multimodal cells of two or all three modalities. The sensory properties of cSTP cells were as follows. 1) Out of 200 cells recorded, 102 (51%) cells were unimodal (59 visual, 33 auditory, and 10 somesthetic), 36 (18%) cells were bimodal (21 visual+auditory, 7 visual+somesthetic, and 8 auditory+somesthetic), and four (2%) cells were trimodal. Visual and auditory responses were more frequent than somesthetic responses: the ratio of the population of cells driven by visual: auditory: somesthetic stimuli was 3:2:1. 2) Visual RFs were large (mean diameter, 59 degrees), but two-thirds were limited to the contralateral visual hemifield. About half the cells showed directional selectivity for moving visual stimuli. None showed selectivity for particular visual shapes.(ABSTRACT TRUNCATED AT 400 WORDS)