Organization of striate cortex of alert, trained monkeys (Macaca fascicularis): ongoing activity, stimulus selectivity, and widths of receptive field activating regions

1995 ◽  
Vol 74 (5) ◽  
pp. 2100-2125 ◽  
Author(s):  
D. M. Snodderly ◽  
M. Gur
Keyword(s):  
Receptive Field ◽  
Macaca Fascicularis ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Cortical Processing ◽  
Ongoing Activity ◽  
Fine Detail ◽  
Laminar Distribution ◽  
Maintained Discharge ◽  
Direction Of Movement

1. In alert macaque monkeys, multiunit activity is encountered in an alternating sequence of silent and spontaneously active zones as an electrode is lowered through the striate cortex (V1). 2. Individual neurons that are spontaneously active in the dark usually have a maintained discharge in the light. Because both types of discharge occur in the absence of deliberate stimulation, we call them the "ongoing" activity. The zones with ongoing activity correspond to the cytochrome oxidase (CytOx)-rich geniculorecipient layers 4A, 4C, and 6, whereas the adjacent layers 2/3, 4B, and 5 have little ongoing activity. 3. The widths of receptive field activating regions (ARs) are positively correlated with the cells' ongoing activity. Cells with larger ARs are preferentially located in the CytOx-rich (input) layers, and many are unselective for stimulus orientation. However, approximately 90% of the cells in the silent layers are orientation selective, and they often have small ARs. 4. The laminar distribution of selectivity for orientation and direction of movement in alert animals is consistent with earlier results from anesthetized animals, but the laminar distribution of AR widths differs. In alert macaques, the ARs of direction-selective cells in layer 4B and of orientation-selective cells in layer 5 are among the smallest in V1. 5. Our findings indicate that the input layers of V1 (4A, 4C, and 6) have a diversity of AR widths, including large ones. Cortical processing produces receptive fields in some of the output layers (4B and 5) that are restricted to small ARs with high resolution of spatial position. These results imply potent lateral and/or interlaminar interactions in alert animals in early cortical processing. The diversity of AR widths generated in V1 may contribute to detection of fine detail in the presence of contrasting backgrounds--the early stages of figure-ground discrimination.

Quantitative studies of single-cell properties in monkey striate cortex. I. Spatiotemporal organization of receptive fields

1976 ◽  
Vol 39 (6) ◽  
pp. 1288-1319 ◽  
Author(s):  
P. H. Schiller ◽  
B. L. Finlay ◽  
S. F. Volman
Keyword(s):  
Receptive Field ◽  
Striate Cortex ◽  
Single Cells ◽  
Receptive Fields ◽  
Spatial Separation ◽  
The Other ◽  
Field Size ◽  
Stimulus Movement ◽  
Direction Of Movement ◽  
Opposite Contrast

1. The properties of single cells in striate cortex of the rhesus monkey, representing the visual field 2 degrees -5 degrees from the fovea, were examined quantitatively with stationary and moving stimuli. Three distinct classes of cells were identified: S type, CX type, and T type. 2. S-type cells were defined as those oriented cells which to the optimal direction of movement in their receptive fields exhibited one or more spatially separate subfields within each of which a response was obtained to either a light or dark edge, but not to both. Several different types of S-cells were distinguished: a) S1-type cells for which moving edges revealed a single excitatory area within which a response was elicited by either a light or a dark edge but not by both. Most of these cells were unidirectional. b) S2-type cells for which moving edges revealed two spatially separate response areas, one of which was excited by a light edge and the other by a dark edge. Both regions responded to the same direction of movement. c) S3-type cells which had two response areas, one of which was excited by a stimulus moving in one direction (at right angles to the axis of orientation) and the other, of opposite contrast, which responded in the opposite direction, d) S4-type cells which to one direction of movement showed two spatially separate regions sensitive to a light and dark edge and which in the other direction of movement had only one responsive area (either light or dark). e) Cells which had multiple spatially separate subfields (S5-7 types). 3. CX-type cells were defined as those oriented cells which in their receptive fields exhibited no spatial separation for light- and dark-edge responses; they discharged to both edges in the same direction of movement and in the same spatial area. Flashing stimuli elicited both on and off responses throughout the receptive field. CX-type cells were predominantly of two types: those which were selective for direction of stimulus movement and those which were not. 4. A third class of cells (T-type) were those which were excited by only one sign of contrast change and responded in a sustained fashion even when there was no contour within the receptive field. These cells were poorly or not at all oriented; some of them were selective to wavelength. 5. Quantitative comparisons showed the following differences between S-type and CX-type cells: a) S-type cells had smaller receptive fields than CX-type cells but the populations over-lapped considerably. Receptive-field size was smallest in layer 4c. In all other layers S-type cells had the same size fields. CX-type cells, by contrast, tended to have larger fields in layer 5-6 than 2-3. b) The spatial separation between light and dark response areas was the best criterion for distinguishing S-type and CX-type cells. The distribution of this measure disclosed two populations of cells with relatively limited overlap. c) In layers 2 and 3, both S-type and CX-type cells had low spontaneous activity…

Ferrier lecture - Functional architecture of macaque monkey visual cortex

1977 ◽  
Vol 198 (1130) ◽  
pp. 1-59 ◽  
Keyword(s):  
Visual Cortex ◽  
Visual Field ◽  
Receptive Field ◽  
Striate Cortex ◽  
Single Cells ◽  
Ocular Dominance ◽  
Receptive Fields ◽  
Macaque Monkey ◽  
Area 17 ◽  
Orientation Specificity

Of the many possible functions of the macaque monkey primary visual cortex (striate cortex, area 17) two are now fairly well understood. First, the incoming information from the lateral geniculate bodies is rearranged so that most cells in the striate cortex respond to specifically oriented line segments, and, second, information originating from the two eyes converges upon single cells. The rearrangement and convergence do not take place immediately, however: in layer IVc, where the bulk of the afferents terminate, virtually all cells have fields with circular symmetry and are strictly monocular, driven from the left eye or from the right, but not both; at subsequent stages, in layers above and below IVc, most cells show orientation specificity, and about half are binocular. In a binocular cell the receptive fields in the two eyes are on corresponding regions in the two retinas and are identical in structure, but one eye is usually more effective than the other in influencing the cell; all shades of ocular dominance are seen. These two functions are strongly reflected in the architecture of the cortex, in that cells with common physiological properties are grouped together in vertically organized systems of columns. In an ocular dominance column all cells respond preferentially to the same eye. By four independent anatomical methods it has been shown that these columns have the form of vertically disposed alternating left-eye and right-eye slabs, which in horizontal section form alternating stripes about 400 μm thick, with occasional bifurcations and blind endings. Cells of like orientation specificity are known from physiological recordings to be similarly grouped in much narrower vertical sheeet-like aggregations, stacked in orderly sequences so that on traversing the cortex tangentially one normally encounters a succession of small shifts in orientation, clockwise or counterclockwise; a 1 mm traverse is usually accompanied by one or several full rotations through 180°, broken at times by reversals in direction of rotation and occasionally by large abrupt shifts. A full complement of columns, of either type, left-plus-right eye or a complete 180° sequence, is termed a hypercolumn. Columns (and hence hypercolumns) have roughly the same width throughout the binocular part of the cortex. The two independent systems of hypercolumns are engrafted upon the well known topographic representation of the visual field. The receptive fields mapped in a vertical penetration through cortex show a scatter in position roughly equal to the average size of the fields themselves, and the area thus covered, the aggregate receptive field, increases with distance from the fovea. A parallel increase is seen in reciprocal magnification (the number of degrees of visual field corresponding to 1 mm of cortex). Over most or all of the striate cortex a movement of 1-2 mm, traversing several hypercolumns, is accompanied by a movement through the visual field about equal in size to the local aggregate receptive field. Thus any 1-2 mm block of cortex contains roughly the machinery needed to subserve an aggregate receptive field. In the cortex the fall-off in detail with which the visual field is analysed, as one moves out from the foveal area, is accompanied not by a reduction in thickness of layers, as is found in the retina, but by a reduction in the area of cortex (and hence the number of columnar units) devoted to a given amount of visual field: unlike the retina, the striate cortex is virtually uniform morphologically but varies in magnification. In most respects the above description fits the newborn monkey just as well as the adult, suggesting that area 17 is largely genetically programmed. The ocular dominance columns, however, are not fully developed at birth, since the geniculate terminals belonging to one eye occupy layer IVc throughout its length, segregating out into separate columns only after about the first 6 weeks, whether or not the animal has visual experience. If one eye is sutured closed during this early period the columns belonging to that eye become shrunken and their companions correspondingly expanded. This would seem to be at least in part the result of interference with normal maturation, though sprouting and retraction of axon terminals are not excluded.

Receptive-field characteristics of neurons in cat striate cortex: Changes with visual field eccentricity

1976 ◽  
Vol 39 (3) ◽  
pp. 512-533 ◽  
Author(s):  
J. R. Wilson ◽  
S. M. Sherman
Keyword(s):  
Visual Field ◽  
Receptive Field ◽  
Striate Cortex ◽  
Ocular Dominance ◽  
Receptive Fields ◽  
Direction Selectivity ◽  
Orientation Selectivity ◽  
Simple Cells ◽  
Complex Cells ◽  
Cat Striate Cortex

1. Receptive-field properties of 214 neurons from cat striate cortex were studied with particular emphasis on: a) classification, b) field size, c) orientation selectivity, d) direction selectivity, e) speed selectivity, and f) ocular dominance. We studied receptive fields located throughtout the visual field, including the monocular segment, to determine how receptivefield properties changed with eccentricity in the visual field.2. We classified 98 cells as "simple," 80 as "complex," 21 as "hypercomplex," and 15 in other categories. The proportion of complex cells relative to simple cells increased monotonically with receptive-field eccenticity.3. Direction selectivity and preferred orientation did not measurably change with eccentricity. Through most of the binocular segment, this was also true for ocular dominance; however, at the edge of the binocular segment, there were more fields dominated by the contralateral eye.4. Cells had larger receptive fields, less orientation selectivity, and higher preferred speeds with increasing eccentricity. However, these changes were considerably more pronounced for complex than for simple cells.5. These data suggest that simple and complex cells analyze different aspects of a visual stimulus, and we provide a hypothesis which suggests that simple cells analyze input typically from one (or a few) geniculate neurons, while complex cells receive input from a larger region of geniculate neurons. On average, this region is invariant with eccentricity and, due to a changing magnification factor, complex fields increase in size with eccentricity much more than do simple cells. For complex cells, computations of this geniculate region transformed to cortical space provide a cortical extent equal to the spread of pyramidal cell basal dendrites.

Spatial properties of neurons in the monkey striate cortex

1987 ◽  
Vol 231 (1263) ◽  
pp. 251-288 ◽  
Keyword(s):  
Receptive Field ◽  
Contrast Sensitivity ◽  
Visual Processing ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Spatial Filter ◽  
Cortical Cells ◽  
Simple Cells ◽  
Gabor Function ◽  
Difference Of Gaussians

Contrast sensitivity as a function of spatial frequency was determined for 138 neurons in the foveal region of primate striate cortex. The accuracy of three models in describing these functions was assessed by the method of least squares. Models based on difference-of-Gaussians (DOG) functions were shown to be superior to those based on the Gabor function or the second differential of a Gaussian. In the most general case of the DOG models, each subregion of a simple cell’s receptive field was constructed from a single DOG function. All the models are compatible with the classical observation that the receptive fields of simple cells are made up of spatially discrete ‘on’ and ‘off’ regions. Although the DOG-based models have more free parameters, they can account better for the variety of shapes of spatial contrast sensitivity functions observed in cortical cells and, unlike other models, they provide a detailed description of the organization of subregions of the receptive field that is consistent with the physiological constraints imposed by earlier stages in the visual pathway. Despite the fact that the DOG-based models have spatially discrete components, the resulting amplitude spectra in the frequency domain describe complex cells just as well as simple cells. The superiority of the DOG-based models as a primary spatial filter is discussed in relation to popular models of visual processing that use the Gabor function or the second differential of a Gaussian.

Functional properties of retinal ganglion cells during optic nerve regeneration in the goldfish

1998 ◽  
Vol 15 (6) ◽  
pp. 1145-1155 ◽  
Author(s):  
D.-J. OH ◽  
D.P.M. NORTHMORE
Keyword(s):  
Receptive Field ◽  
Ganglion Cells ◽  
Receptive Fields ◽  
Optic Tract ◽  
Evoked Responses ◽  
Antidromic Activation ◽  
Discharge Rates ◽  
Conduction Velocities ◽  
Maintained Discharge ◽  
Retinotectal System

After being severed, optic axons in goldfish regenerate and eventually restore the retinotectal map; refinement of the map depends upon impulse activity generated by the ganglion cells. Because little is known about the changes in activity and receptive-field properties of ganglion cells during regeneration, we made extracellular recordings from them in the intact eye up to 95 days after sectioning their axons in the optic tract. Their receptive fields were classified as OFF-, ON–OFF-, or ON-centers, and their axonal conduction velocities measured by antidromic activation. The rate of encountering single units dropped drastically at 4–8 days postsection when only a few OFF-center units could be recorded, recovering to normal between 42 and 63 days. Receptive-field centers were normal in size, except for the few OFF-centers at 4–8 days which were abnormally large. Maintained discharge rates of all types were depressed up to 42 days, but ON–OFF-center units were more spontaneously active than normal around 42 days. Light-evoked responses in OFF-center units were subnormal at 4–8 days, becoming supernormal at 16 days and normal thereafter. ON–OFF- and ON-center units started to regain responsiveness at 16 days, and became supernormal at 42 days, before returning to normal. Conduction velocities of all fiber groups dropped to a minimum at 8 days, the fastest being affected most. There was a gradual recovery to normal conduction velocity by 63 days. The conduction latencies of OFF- and ON–OFF-center units recovered to normal by 42 days, and ON-center units by 63 days. Recovery of ganglion cell responsiveness correlates with functional recovery in the retinotectal system: OFF-center units recover light-evoked responses at about the time OFF activity first reappears in the tectum. ON- and ON–OFF-center units recover later, exhibiting supernormal spiking activity around the time that ON responses reappear in the tectum.

Receptive-field properties of neurons in different laminae of visual cortex of the cat

1978 ◽  
Vol 41 (4) ◽  
pp. 948-962 ◽  
Author(s):  
A. G. Leventhal ◽  
H. V. Hirsch
Keyword(s):  
Visual Cortex ◽  
Receptive Field ◽  
Striate Cortex ◽  
Visual Stimulation ◽  
Receptive Fields ◽  
Spontaneous Discharge ◽  
C Cells ◽  
Receptive Field Properties ◽  
Discharge Rates ◽  
Y Cells

1. Receptive-field properties of neurons in the different layers of the visual cortex of normal adult cats were analyzed quantitatively. Neurons were classified into one of two groups: 1) S-cells, which have discrete on- and/or off-regions in their receptive fields and possess inhibitory side bands; 2) C-cells, which do not have discrete on- and off-regions in their receptive fields but display an on-off response to flashing stimuli. Neurons of this type rarely display side-band inhibition. 2. As a group, S-cells display lower relative degrees of binocularity and are more selective for stimulus orientation than C-cells. In addition, within a given lamina the S-cells have smaller receptive fields, lower cutoff velocities, lower peak responses to visual stimulation, and lower spontaneous activity than do the C-cells. 3. S-cells in all layers of the cortex display similar orientation sensitivities, mean spontaneous discharge rates, peak response to visual stimulation, and degrees of binocularity. 4. Many of the receptive-field properties of cortical cells vary with laminar location. Receptive-field sizes and cutoff velocities of S-cells and of C-cells are greater in layers V and VI than in layers II-IV. For S-cells, preferred velocities are also greater in layers V and VI than in layers II-IV. Furthermore, C-cells in layers V and VI display high mean spontaneous discharge rates, weak orientation preferences, high relative degrees of binocularity, and higher peak responses to visual stimulation when compared to C-cells in layers II and III. 5. The receptive-field properties of cells in layers V-VI of the striate cortex suggest that most neurons that have their somata in these laminae receive afferents from LGNd Y-cells. Hence, our results suggest that afferents from LGNd Y-cells may play a major part in the cortical control of subcortical visual functions.

scholarly journals The two-dimensional spatial structure of simple receptive fields in cat striate cortex

1987 ◽  
Vol 58 (6) ◽  
pp. 1187-1211 ◽  
Author(s):  
J. P. Jones ◽  
L. A. Palmer
Keyword(s):  
Receptive Field ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Correlation Method ◽  
Reverse Correlation ◽  
Moving Stimuli ◽  
Response Profile ◽  
Spatial Dimensions ◽  
Cat Striate Cortex

1. A reverse correlation (6, 8, 25, 35) method is developed that allows quantitative determination of visual receptive-field structure in two spatial dimensions. This method is applied to simple cells in the cat striate cortex. 2. It is demonstrated that the reverse correlation method yields results with several desirable properties, including convergence and reproducibility independent of modest changes in stimulus parameters. 3. In contrast to results obtained with moving stimuli, we find that the bright and dark excitatory subregions in simple receptive fields do not overlap to any great extent. This difference in results may be attributed to confounding the independent variables space and time when using moving stimuli. 4. All simple receptive fields have subregions that vary smoothly in all directions in space. There are no sharp transitions either between excitatory subregions or between subregions and the area surrounding the receptive field. 5. Simple receptive fields vary both in the number of subregions observed, in the elongation of each subregion, and in the overall elongation of the field. In contrast with results obtained using moving stimuli, we find that subregions within a given receptive field need not be the same length. 6. The hypothesis that simple receptive fields can be modeled as either even symmetric or odd symmetric about a central axis is evaluated. This hypothesis is found to be false in general. Most simple receptive fields are neither even symmetric nor odd symmetric. 7. The hypothesis that simple receptive fields can be modeled as the product of a width response profile and an orthogonal length response profile (Cartesian separability) is evaluated. This hypothesis is found to be true for only approximately 50% of the cells in our sample.

Development of spatial receptive-field organization and orientation selectivity in kitten striate cortex

1985 ◽  
Vol 53 (5) ◽  
pp. 1158-1178 ◽  
Author(s):  
B. O. Braastad ◽  
P. Heggelund
Keyword(s):  
Receptive Field ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Orientation Selectivity ◽  
Half Width ◽  
Orientation Tuning ◽  
Maximal Response ◽  
Tuning Curves ◽  
Field Organization ◽  
Receptive Field Organization

The functional organization of the receptive field of neurons in striate cortex of kittens from 8 days to 3 mo of age was studied by extracellular recordings. A quantitative dual-stimulus technique was used, which allowed for analysis of both enhancement and suppression zones in the receptive field. Furthermore the development of orientation selectivity was studied quantitatively in the same cells. Already in the youngest kittens the receptive fields were spatially organized like adult fields, with a central zone and adjacent flanks that responded in opposite manner to the light stimulus. The relative suppression in the subzones was as strong as in adult cells. Both simple and complex cells were found from 8 days. The receptive fields were like magnified adult fields. The width of the dominant discharge-field zone and the distance between the positions giving maximum discharge and maximum suppression decreased with age in the same proportions. The decrease could be explained by a corresponding decrease of the receptive-field-center size of retinal ganglion cells. Forty percent of the cells were orientation selective before 2 wk, and the fraction increased to 94% at 4 wk. Cells whose responses could be attenuated to at least half of the maximal response by changes of slit orientation were termed orientation selective. The half-width of the orientation-tuning curves narrowed during the first 5 wk, and this change was most marked in simple cells. The ability of the cells to discriminate between orientations in statistical terms was weak in the youngest kittens due to a large response variability, and showed a more pronounced development than the half-width did. The orientation-tuning curves were fitted by an exponential function, which showed the shape to be adultlike in all age groups. Two kittens were dark reared until recording at 1 mo of age. The spatial receptive-field organization and the orientation selectivity in these kittens were similar to normal-reared kittens at 1 mo. The responsivity of the cells of the dark-reared kittens was lower, and the latency before firing was longer than in the normal-reared kittens of the same age, and these response properties were more similar to those in 1- to 2-wk-old normal kittens. Our results indicate that the spatial organization of the receptive field is innate in most cells and that visual experience is unnecessary for the organization to be maintained and for the receptive-field width to mature during the first month postnatally.(ABSTRACT TRUNCATED AT 400 WORDS)

Spatiotemporal organization of simple-cell receptive fields in the cat's striate cortex. I. General characteristics and postnatal development

1993 ◽  
Vol 69 (4) ◽  
pp. 1091-1117 ◽  
Author(s):  
G. C. DeAngelis ◽  
I. Ohzawa ◽  
R. D. Freeman
Keyword(s):  
Receptive Field ◽  
Postnatal Development ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Space Time ◽  
Field Structure ◽  
Temporal Response ◽  
Spatial Frequencies ◽  
Spatiotemporal Receptive Field ◽  
Time Courses

1. Most studies of cortical neurons have focused on the spatial structure of receptive fields. For a more complete functional description of these neurons, it is necessary to consider receptive-field structure in the joint domain of space and time. We have studied the spatiotemporal receptive-field structure of 233 simple cells recorded from the striate cortex of adult cats and kittens at 4 and 8 wk postnatal. The dual goal of this study is to provide a detailed quantitative description of spatiotemporal receptive-field structure and to compare the developmental time courses of spatial and temporal response properties. 2. Spatiotemporal receptive-field profiles have been measured with the use of a reverse correlation method, in which we compute the cross-correlation between a neuron's response and a random sequence of small, briefly presented bright and dark stimuli. The receptive-field profiles of some simple cells are space-time separable, meaning that spatial and temporal response characteristics can be dissociated. Other cells have receptive-field profiles that are space-time inseparable. In these cases, a particular spatial location cannot be designated, unambiguously, as belonging to either an on or off subregion. However, separate on and off subregions may be clearly distinguished in the joint space-time domain. These subregions are generally tilted along an oblique axis. 3. Our observations show that spatial and temporal aspects of receptive-field structure mature with clearly different time courses. By 4 wk postnatal, the spatial symmetry and periodicity of simple-cell receptive fields have reached maturity. The spatial extent (or size) of these receptive fields is adult-like by 8 wk postnatal. In contrast, the response latency and time duration of spatiotemporal receptive fields do not mature until well beyond 8 wk postnatal. 4. By applying Fourier analysis to spatiotemporal receptive-field profiles, we have examined the postnatal development of spatial and temporal selectivity in the frequency domain. By 8 wk postnatal, spatial frequency tuning has clearly reached maturity. On the contrary, temporal frequency selectivity remains markedly immature at 8 wk. We have also examined the joint distribution of optimal spatial and temporal frequencies. From 4 wk postnatal until 8 wk postnatal, the range of optimal spatial frequencies increases substantially, whereas the range of optimal temporal frequencies remains largely unchanged. From 8 wk postnatal until adulthood, there is a large increase in optimal temporal frequencies for cells tuned to low spatial frequencies. For cells tuned to high spatial frequencies, the distribution of optimal temporal frequencies does not change much beyond 8 wk postnatal.(ABSTRACT TRUNCATED AT 400 WORDS)

scholarly journals Local Signals From Beyond the Receptive Fields of Striate Cortical Neurons

2003 ◽  
Vol 90 (2) ◽  
pp. 822-831 ◽  
Author(s):  
James R. Müller ◽  
Andrew B. Metha ◽  
John Krauskopf ◽  
Peter Lennie
Keyword(s):  
Receptive Field ◽  
Cortical Neurons ◽  
Striate Cortex ◽  
Image Representation ◽  
Receptive Fields ◽  
Preferred Orientation ◽  
Orientation Tuning ◽  
Stimulus Space ◽  
Tuning Curves ◽  
Complex Cells

We examined in anesthetized macaque how the responses of a striate cortical neuron to patterns inside the receptive field were altered by surrounding patterns outside it. The changes in a neuron's response brought about by a surround are immediate and transient: they arise with the same latency as the response to a stimulus within the receptive field (this argues for a source locally in striate cortex) and become less effective as soon as 27 ms later. Surround signals appeared to exert their influence through divisive interaction (normalization) with those arising in the receptive field. Surrounding patterns presented at orientations slightly oblique to the preferred orientation consistently deformed orientation tuning curves of complex (but not simple) cells, repelling the preferred orientation but without decreasing the discriminability of the preferred grating and ones at slightly oblique orientations. By reducing responsivity and changing the tuning of complex cells locally in stimulus space, surrounding patterns reduce the correlations among responses of neurons to a particular stimulus, thus reducing the redundancy of image representation.

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