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 Strobe Rearing Reduces Direction Selectivity in Area 17 by Altering Spatiotemporal Receptive-Field Structure

1998 ◽  
Vol 80 (6) ◽  
pp. 2991-3004 ◽  
Author(s):  
Allen L. Humphrey ◽  
Alan B. Saul
Keyword(s):  
Receptive Field ◽  
Frequency Tuning ◽  
Temporal Frequency ◽  
Receptive Fields ◽  
Direction Selectivity ◽  
Field Structure ◽  
Area 17 ◽  
Directional Tuning ◽  
Simple Cells ◽  
Spatiotemporal Receptive Field

Humphrey, Allen L. and Alan B. Saul. Strobe rearing reduces direction selectivity in area 17 by altering spatiotemporal receptive-field structure. J. Neurophysiol. 80: 2991–3004, 1998. Direction selectivity in simple cells of cat area 17 is linked to spatiotemporal (S-T) receptive-field structure. S-T inseparable receptive fields display gradients of response timing across the receptive field that confer a preferred direction of motion. Receptive fields that are not direction selective lack gradients; they are S-T separable, displaying uniform timing across the field. Here we further examine this link using a developmental paradigm that disrupts direction selectivity. Cats were reared from birth to 8 mo of age in 8-Hz stroboscopic illumination. Direction selectivity in simple cells was then measured using gratings drifting at different temporal frequencies (0.25–16 Hz). S-T structure was assessed using stationary bars presented at different receptive-field positions, with bar luminance being modulated sinusoidally at different temporal frequencies. For each cell, plots of response phase versus bar position were fit by lines to characterize S-T inseparability at each temporal frequency. Strobe rearing produced a profound loss of direction selectivity at all temporal frequencies; only 10% of cells were selective compared with 80% in normal cats. The few remaining directional cells were selective over a narrower than normal range of temporal frequencies and exhibited weaker than normal direction selectivity. Importantly, the directional loss was accompanied by a virtual elimination of S-T inseparability. Nearly all cells were S-T separable, like nondirectional cells in normal cats. The loss was clearest in layer 4. Normally, inseparability is greatest there, and it correlates well ( r = 0.77) with direction selectivity; strobe rearing reduced inseparability and direction selectivity to very low values. The few remaining directional cells were inseparable. In layer 6 of normal cats, most direction-selective cells are only weakly inseparable, and there is no consistent relationship between the two measures. However, after strobe rearing, even the weak inseparability was eliminated along with direction selectivity. The correlated changes in S-T structure and direction selectivity were confirmed using conventional linear predictions of directional tuning based on responses to counterphasing bars and white noise stimuli. The developmental changes were permanent, being observed up to 12 yr after strobe rearing. The deficits were remarkably specific; strobe rearing did not affect spatial receptive-field structure, orientation selectivity, spatial or temporal frequency tuning, or general responsiveness to visual stimuli. These results provide further support for a critical role of S-T structure in determining direction selectivity in simple cells. Strobe rearing eliminates directional tuning by altering the timing of responses within the receptive field.

Inhibitory Contributions to Spatiotemporal Receptive-Field Structure and Direction Selectivity in Simple Cells of Cat Area 17

1999 ◽  
Vol 81 (3) ◽  
pp. 1212-1224 ◽  
Author(s):  
Aditya Murthy ◽  
Allen L. Humphrey
Keyword(s):  
Receptive Field ◽  
Receptive Fields ◽  
Direction Selectivity ◽  
Field Structure ◽  
Area 17 ◽  
Directional Tuning ◽  
Simple Cells ◽  
Layer 4 ◽  
Two Measures ◽  
Spatiotemporal Receptive Field

Inhibitory contributions to spatiotemporal receptive-field structure and direction selectivity in simple cells of cat area 17. Intracortical inhibition contributes to direction selectivity in primary visual cortex, but how it acts has been unclear. We investigated this problem in simple cells of cat area 17 by taking advantage of the link between spatiotemporal (S-T) receptive-field structure and direction selectivity. Most cells in layer 4 have S-T–oriented receptive fields in which gradients of response timing across the field confer a preferred direction of motion. Linear summation of responses across the receptive field, followed by a static nonlinear amplification, has been shown previously to account for directional tuning in layer 4. We tested the hypotheses that inhibition acts by altering S-T structure or the static nonlinearity or both. Drifting and counterphasing sinewave gratings were used to measure direction selectivity and S-T structure, respectively, in 17 layer 4 simple cells before and during iontophoresis of bicuculline methiodide (BMI), a GABAA antagonist. S-T orientation was quantified from fits to response temporal phase versus stimulus spatial phase data. Bicuculline reduced direction selectivity and S-T orientation in nearly all cells, and reductions in the two measures were well correlated ( r = 0.81) and reversible. Using conventional linear predictions based on response phase and amplitude, we found that BMI-induced changes in S-T structure also accounted well for absolute changes in the amplitude and phase of responses to gratings drifting in the preferred and nonpreferred direction. For each cell we also calculated an exponent used to estimate the static nonlinearity. Bicuculline reduced the exponent in most cells, but the changes were not correlated with reductions in direction selectivity. We conclude that GABAA-mediated inhibition influences directional tuning in layer 4 primarily by sculpting S-T receptive-field structure. The source of the inhibition is likely to be other simple cells with certain spatiotemporal relationships to their target. Despite reductions in the two measures, most receptive fields maintained some directional tuning and S-T orientation during BMI. This suggests that their excitatory inputs, arising from the lateral geniculate nucleus and within area 17, are sufficient to create some S-T orientation and that inhibition accentuates it. Finally, BMI also reduced direction selectivity in 8 of 10 simple cells tested in layer 6, but the reductions were not accompanied by systematic changes in S-T structure. This reflects the fact that S-T orientation, as revealed by our first-order measures of the receptive field, is weak there normally. Inhibition likely affects layer 6 cells via more complex, nonlinear interactions.

Spatiotemporal Receptive Field Organization in the Lateral Geniculate Nucleus of Cats and Kittens

1997 ◽  
Vol 78 (2) ◽  
pp. 1045-1061 ◽  
Author(s):  
Daqing Cai ◽  
Gregory C. Deangelis ◽  
Ralph D. Freeman
Keyword(s):  
Receptive Field ◽  
Lateral Geniculate Nucleus ◽  
Spatial Organization ◽  
Field Structure ◽  
Temporal Response ◽  
Temporal Phase ◽  
Lateral Geniculate ◽  
Spatiotemporal Receptive Field ◽  
Field Organization ◽  
Receptive Field Organization

Cai, Daqing, Gregory C. DeAngelis, and Ralph D. Freeman. Spatiotemporal receptive field organization in the lateral geniculate nucleus of cats and kittens. J. Neurophysiol. 78: 1045–1061, 1997. We have studied the spatiotemporal receptive-field organization of 144 neurons recorded from the dorsal lateral geniculate nucleus (dLGN) of adult cats and kittens at 4 and 8 wk postnatal. Receptive-field profiles were obtained with the use of a reverse correlation technique, in which we compute the cross-correlation between the action potential train of a neuron and a randomized sequence of long bright and dark bar stimuli that are flashed throughout the receptive field. Spatiotemporal receptive-field profiles of LGN neurons generally exhibit a biphasic temporal response, as well as the classical center-surround spatial organization. For nonlagged cells, the first temporal phase of the response dominates, whereas for lagged neurons, the second temporal phase of the response is typically the largest. This temporal phase difference between lagged and nonlagged cells accounts for their divergent behavior in response to flashed stimuli. Most LGN cells exhibit some degree of space-time inseparability, which means that the receptive field cannot simply be viewed as the product of a spatial waveform and a temporal waveform. In these cases, the response of the surround is typically delayed relative to that of the center, and there is some blending of center and surround during the time course of the response. We demonstrate that a simple extension of the traditional difference-of-Gaussians (DOG) model, in which the surround response is delayed relative to that of the center, accounts nicely for these findings. With regard to development, our analysis shows that spatial and temporal aspects of receptive field structure mature with markedly different time courses. After 4 wk postnatal, there is little change in the spatial organization of LGN receptive fields, with the exception of a weak, but significant, trend for the surround to become smaller and stronger with age. In contrast, there are substantial changes in temporal receptive-field structure after 4 wk postnatal. From 4 to 8 wk postnatal, the shape of the temporal response profile changes, becoming more biphasic, but the latency and duration of the response remain unchanged. From 8 wk postnatal to adulthood, the shape of the temporal profile remains approximately constant, but there is a dramatic decline in both the latency and duration of the response. Comparison of our results with recent data from cortical (area 17) simple cells reveals that the temporal development of LGN cells accounts for a substantial portion of the temporal maturation of simple cells.

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.

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.

Effects of strabismus on responsivity, spatial resolution, and contrast sensitivity of cat lateral geniculate neurons

1984 ◽  
Vol 52 (3) ◽  
pp. 538-552 ◽  
Author(s):  
K. R. Jones ◽  
R. E. Kalil ◽  
P. D. Spear
Keyword(s):  
Contrast Sensitivity ◽  
Spatial Resolution ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Orienting Response ◽  
Lateral Geniculate ◽  
Spatial Frequencies ◽  
Area Centralis ◽  
X Cells ◽  
Y Cells

Rearing cats with esotropia is known to cause a number of deficits in visual behavior tested through the deviated eye. These include a loss of orienting response to stimuli presented in the nasal visual field of the deviated eye, a reduction in visual acuity, and a general reduction in contrast sensitivity at all spatial frequencies. To assess the involvement of the lateral geniculate nucleus (LGN) in these deficits, we measured the following: 1) the visual responsiveness of lamina A1 cells with peripheral (more than 10 degrees from area centralis) receptive fields in three esotropic and three normal cats and 2) the spatial resolution and contrast sensitivity of lamina A X-cells with central (within 5 degrees of the area centralis) receptive fields in six esotropic and six normal cats. For comparison, we also measured LGN X-cell spatial resolutions in four exotropic cats and in two cats raised with an esotropia in one eye and the lids of the other eye sutured shut (MD-estropes). Recordings from the lateral portion of lamina A1 in esotropic cats yielded similar numbers of visually responsive cells with far nasal receptive fields as were seen in normal animals. Peak and mean response rates to a flashing spot also were normal. In addition, no differences were found between esotropes and normals in the percentages of X- and Y-cells encountered. These results suggest that the loss of orienting response to stimuli presented in the nasal field (12, 20) is not due to a loss of neural responses in the LGN of esotropic cats. In addition, they suggest that decreases in cell size in lamina A1 of esotropic cats (13, 36; R. E. Kalil, unpublished observations) are not accompanied by marked functional abnormalities of the cells and that cortical abnormalities ipsilateral to the deviated eye (22) are likely to have their origin within striate cortex itself. Recordings from lamina A cells with receptive fields near area centralis revealed that the average X-cell spatial resolution in esotropes (2.1 cycles/deg) was significantly lower than that in normal cats (3.1 cycles/deg). This reduction was seen in all esotropic cats tested and was due both to an increase in the proportion of X-cells with very low spatial resolution and to a loss of X-cells responding to high spatial frequencies (greater than 3.25 cycles/deg). The average spatial resolution of X-cells driven by the deviated eye in MD-esotropes fell midway between those of esotropes and normals. In exotropes, mean X-cell spatial resolution was normal.(ABSTRACT TRUNCATED AT 400 WORDS)

scholarly journals Attention operates uniformly throughout the classical receptive field and the surround

eLife ◽  
2016 ◽  
Vol 5 ◽  
Author(s):  
Bram-Ernst Verhoef ◽  
John HR Maunsell
Keyword(s):  
Receptive Field ◽  
Cortical Neurons ◽  
Receptive Fields ◽  
Spatial Summation ◽  
Field Structure ◽  
Neuronal Responses ◽  
Large Variability ◽  
Area V4 ◽  
Classical Receptive Field ◽  
Spatially Varying

Shifting attention among visual stimuli at different locations modulates neuronal responses in heterogeneous ways, depending on where those stimuli lie within the receptive fields of neurons. Yet how attention interacts with the receptive-field structure of cortical neurons remains unclear. We measured neuronal responses in area V4 while monkeys shifted their attention among stimuli placed in different locations within and around neuronal receptive fields. We found that attention interacts uniformly with the spatially-varying excitation and suppression associated with the receptive field. This interaction explained the large variability in attention modulation across neurons, and a non-additive relationship among stimulus selectivity, stimulus-induced suppression and attention modulation that has not been previously described. A spatially-tuned normalization model precisely accounted for all observed attention modulations and for the spatial summation properties of neurons. These results provide a unified account of spatial summation and attention-related modulation across both the classical receptive field and the surround.

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.

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