Temporal encoding of two-dimensional patterns by single units in primate primary visual cortex. I. Stimulus-response relations

1990 ◽  
Vol 64 (2) ◽  
pp. 351-369 ◽  
Author(s):  
B. J. Richmond ◽  
L. M. Optican ◽  
H. Spitzer
Keyword(s):  
Visual Processing ◽  
Cortical Neurons ◽  
Two Dimensional ◽  
Temporal Modulation ◽  
Gray Background ◽  
Complex Cells ◽  
Stimulus Response ◽  
Black And White ◽  
Stimulus Set ◽  
Simple And Complex Cells

1. Previously we developed a new approach for investigating visual system neuronal activity in which single neurons are considered to be communication channels transmitting stimulus-dependent codes in their responses. Application of this approach to the stimulus-response relations of inferior temporal (IT) neurons showed that these carry stimulus-dependent information in the temporal modulation as well as in the strength of their responses. IT cortex is a late station in the visual processing stream. Presumably the neuronal properties arise from the properties of the inputs. However, the discovery that IT neuronal spike trains transmit information in stimulus-dependent temporally modulated codes could not be assumed to be true for those earlier stations, so the techniques used in the earlier study were applied to single-striate cortical neurons in the studies reported here. 2. Single-striate cortical neurons were recorded from three awake, fixating rhesus monkeys. The neurons were stimulated by two sets of patterns. The first set was made up of 128 black-and-white patterns based on a complete, orthogonal set of two-dimensional Walsh-Hadamard functions. These stimuli appear as combinations of black-and-white rectangles and squares, and they fully span the range of all possible black-and-white pictures that can be constructed in an 8 x 8 grid. Except for the stimulus that appeared as an all-white or all-black square, each stimulus had equal areas of white and black. The second stimulus set was made up of single bars constructed in the same 8 x 8 grid as the Walsh stimuli. These were presented both as black against a gray background and white against a gray background. The stimuli were centered on the receptive field, and each member of the stimulus set was presented once before any stimulus appeared again. 3. The responses of 21 striate cortical neurons were recorded and analyzed. Two were identified as simple cells and the other 19 as complex cells according to the criteria originally used by Hubel and Wiesel. The stimulus set elicited a wide variety of response strengths and patterns from each neuron. The responses from both the bars and the Walsh set could be used to differentiate and classify simple and complex cells. 4. The responses of both simple and complex cells showed striking stimulus-related strength and temporal modulation. For all of the complex cells there were instances where the responses to a stimulus and its contrast-reversed mate were substantially different in response strength or pattern, or both.(ABSTRACT TRUNCATED AT 400 WORDS)

Interactive effects among several stimulus parameters on the responses of striate cortical complex cells

1991 ◽  
Vol 66 (2) ◽  
pp. 379-389 ◽  
Author(s):  
T. J. Gawne ◽  
B. J. Richmond ◽  
L. M. Optican
Keyword(s):  
Striate Cortex ◽  
Neuronal Response ◽  
Receptive Fields ◽  
Response Strength ◽  
Interactive Effects ◽  
Temporal Modulation ◽  
Complex Cells ◽  
Black And White ◽  
Interaction Terms ◽  
Stimulus Parameters

1. Although neurons within the visual system are often described in terms of their responses to particular patterns such as bars and edges, they are actually sensitive to many different stimulus features, such as the luminances making up the patterns and the duration of presentation. Many different combinations of stimulus parameters can result in the same neuronal response, raising the problem of how the nervous system can extract information about visual stimuli from such inherently ambiguous responses. It has been shown that complex cells transmit significant amounts of information in the temporal modulation of their responses, raising the possibility that different stimulus parameters are encoded in different aspects of the response. To find out how much information is actually available about individual stimulus parameters, we examined the interactions among three stimulus parameters in the temporally modulated responses of striate cortical complex cells. 2. Sixteen black and white patterns were presented to two awake monkeys at each of four luminance-combinations and five durations, giving a total of 320 unique stimuli. Complex cells were recorded in layers 2 and 3 of striate cortex, with the stimuli centered on the receptive fields as determined by mapping with black and white bars. 3. An analysis of variance (ANOVA) was applied to these data with the three stimulus parameters of pattern, the luminance-combinations, and duration as the independent variables. The ANOVA was repeated with the magnitude and three different aspects of the temporal modulation of the response as the dependent variables. For the 19 neurons studied, many of the interactions between the different stimulus parameters were statistically significant. For some response measures the interactions accounted for more than one-half of the total response variance. 4. We also analyzed the stimulus-response relationships with the use of information theoretical techniques. We defined input codes on the basis of each stimulus parameter alone, as well as their combinations, and output codes on the basis of response strength, and on three measures of temporal modulation, also taken individually and together. Transmitted information was greatest when the response of a neuron was interpreted as a temporally modulated message about combinations of all three stimulus parameters. The interaction terms of the ANOVA suggest that the response of a complex cell can only be interpreted as a message about combinations of all three stimulus parameters.(ABSTRACT TRUNCATED AT 400 WORDS)

Temporal encoding of two-dimensional patterns by single units in primate primary visual cortex. II. Information transmission

1990 ◽  
Vol 64 (2) ◽  
pp. 370-380 ◽  
Author(s):  
B. J. Richmond ◽  
L. M. Optican
Keyword(s):  
Cortical Neurons ◽  
Activity Patterns ◽  
Temporal Cortex ◽  
Principal Component ◽  
Response Strength ◽  
Inferior Temporal Cortex ◽  
Temporal Code ◽  
Response Code ◽  
Stimulus Response ◽  
Black And White

1. Previously, we studied how picture information was processed by neurons in inferior temporal cortex. We found that responses varying in both response strength and temporal waveform carried information about briefly flashed stationary black-and-white patterns. Now, we have applied that same paradigm to the study of striate cortical neurons. 2. In this approach the responses to a set of basic black and white pictures were quantified through use of a set of basic waveforms, the principal components (extracted from all the responses of each neuron). We found that the first principal component, which corresponds to the response strength, and others, which correspond to different basic temporal activity patterns, were significantly related to the stimuli, i.e., the stimulus drove both the response strength and its temporal pattern. 3. Our previous study had shown that, when information theory was used to quantify the stimulus-response relation, inferior temporal neurons convey over twice as much information in a response code that includes temporal modulation as in a response code that includes only the response strength. This study shows that striate cortical neurons also carry twice as much information in a temporal code as in a response strength code. Thus single visual neurons at both ends of a cortical processing chain for visual pattern use a multidimensional temporal code to carry stimulus-related information. 4. These results support our multiplex-filter hypothesis, which states that single visual system neurons can be regarded as several simultaneously active parallel channels, each of which conveys independent information about the stimulus.

The time course of direction-selective adaptation in simple and complex cells in cat striate cortex

1993 ◽  
Vol 70 (5) ◽  
pp. 2024-2034 ◽  
Author(s):  
D. Giaschi ◽  
R. Douglas ◽  
S. Marlin ◽  
M. Cynader
Keyword(s):  
Time Constant ◽  
Cortical Neurons ◽  
Time Course ◽  
Adaptation Period ◽  
Simple Cells ◽  
Complex Cells ◽  
Prolonged Stimulation ◽  
Preferred Direction ◽  
Simple And Complex Cells ◽  
Exponential Change

1. Responses of single cortical neurons in area 17 of anesthetized cats were recorded in response to prolonged stimulation with a patch of drifting square-wave grating. 2. During adaptation in the preferred direction, all neurons showed some reduction in response to motion in the stimulated direction and most showed some reduction in the opposite, nonstimulated direction. 3. For complex cells, the time course of response decrement in both the stimulated and nonstimulated directions was exponential, with an average time constant of 5 s. Response recovery was also exponential but significantly slower, with time constants of 8 and 13 s in the stimulated and nonstimulated directions, respectively. 4. For simple cells the dynamics of the adaptation effect depended on the direction of testing. In the nonstimulated direction the time course of the change in sensitivity was similar to that of complex cells. In the stimulated direction during both the adaptation and recovery periods, simple cells showed an initial rapid exponential change on the order of a few seconds that was followed by a more gradual exponential change. 5. During prolonged stimulation in the nonpreferred direction, there was less overall change in sensitivity. For some neurons the change in sensitivity during adaptation and recovery was exponential, with a short time constant for both simple and complex cells and for stimulated and nonstimulated directions. Other neurons showed no change in sensitivity in either direction and a few neurons showed facilitation during the adaptation period. 6. There appears to be a rapid general or nonspecific process, which may be related to contrast gain control, underlying motion adaptation in striate cortical neurons. An additional slow, direction-selective process is revealed when simple but not complex cells are stimulated in the preferred direction. We suggest that this latter type of adaptation is a key feature underlying the perceptual motion aftereffect.

Direction-selective adaptation in simple and complex cells in cat striate cortex

1988 ◽  
Vol 59 (4) ◽  
pp. 1314-1330 ◽  
Author(s):  
S. G. Marlin ◽  
S. J. Hasan ◽  
M. S. Cynader
Keyword(s):  
Cortical Neurons ◽  
Striate Cortex ◽  
Selective Adaptation ◽  
Direction Selectivity ◽  
Simple Cells ◽  
Complex Cells ◽  
Preferred Direction ◽  
Directional Preference ◽  
Simple And Complex Cells ◽  
Cat Striate Cortex

1. The selectivity of adaptation to unidirectional motion was examined in neurons of the cat striate cortex. Following prolonged stimulation with a unidirectional high-contrast grating, the responsivity of cortical neurons was reduced. In many units this decrease was restricted to the direction of prior stimulation. This selective adaptation produced changes in the degree of direction selectivity of the cortical units (as measured by the ratio of the response to motion in the preferred direction to that in the nonpreferred direction). 2. The initial strength of the directional preference of a given cortical unit did not determine the degree of direction-selective adaptation. Indeed, even non-direction-selective units could exhibit pronounced direction-selective adaptation. The degree of direction-selective adaptation was also independent of the overall decrease in responsivity during adaptation. 3. There was no difference between simple and complex cells in the total amount of adaptation observed. The selectivity of the adaptation, however, did differ between these two cell types. As a group, simple cells showed significant direction-selective adaptation, whereas complex cells did not. The directional preference of most simple cells decreased following preferred direction adaptation and many highly direction selective simple cells became non-direction selective. In addition, simple cells became significantly more direction selective following nonpreferred direction adaptation. 4. Some complex cells also demonstrated direction-selective adaptation. There was, however, much more variability among complex cells than simple cells. Some complex cells actually increased direction selectivity following preferred direction adaptation. These differences between simple and complex cells suggest that changes in direction selectivity following unidirectional adaptation are not due to simple neuronal fatigue of the unit being recorded, but depend on selective adaptation of afferent inputs to the unit. 5. The spontaneous activity of many cortical neurons decreased following preferred direction adaptation but increased following adaptation in the nonpreferred direction. The response to a stationary grating also decreased following preferred direction adaptation. However, there was very little change in the response to a stationary grating following adaptation in the nonpreferred direction.

Two-dimensional receptive-field organization in striate cortical neurons of the cat

1994 ◽  
Vol 11 (4) ◽  
pp. 703-720 ◽  
Author(s):  
Ming Sun ◽  
A. B. Bonds
Keyword(s):  
Receptive Field ◽  
Cortical Neurons ◽  
Ocular Dominance ◽  
Receptive Fields ◽  
Field Size ◽  
Two Dimensional ◽  
Cortical Layers ◽  
Field Organization ◽  
Receptive Field Organization ◽  
Cortical Receptive Fields

AbstractThe two-dimensional organization of receptive fields (RFs) of 44 cells in the cat visual cortex and four cells from the cat LGN was measured by stimulation with either dots or bars of light. The light bars were presented in different positions and orientations centered on the RFs. The RFs found were arbitrarily divided into four general types: Punctate, resembling DOG filters (11%); those resembling Gabor filters (9%); elongate (36%); and multipeaked-type (44%). Elongate RFs, usually found in simple cells, could show more than one excitatory band or bifurcation of excitatory regions. Although regions inhibitory to a given stimulus transition (e.g. ON) often coincided with regions excitatory to the opposite transition (e.g. OFF), this was by no means the rule. Measurements were highly repeatable and stable over periods of at least 1 h. A comparison between measurements made with dots and with bars showed reasonable matches in about 40% of the cases. In general, bar-based measurements revealed larger RFs with more structure, especially with respect to inhibitory regions. Inactivation of lower cortical layers (V-VI) by local GABA injection was found to reduce sharpness of detail and to increase both receptive-field size and noise in upper layer cells, suggesting vertically organized RF mechanisms. Across the population, some cells bore close resemblance to theoretically proposed filters, while others had a complexity that was clearly not generalizable, to the extent that they seemed more suited to detection of specific structures. We would speculate that the broadly varying forms of cat cortical receptive fields result from developmental processes akin to those that form ocular-dominance columns, but on a smaller scale.

Binocular Combination of Contrast Signals by Striate Cortical Neurons in the Monkey

1997 ◽  
Vol 78 (1) ◽  
pp. 366-382 ◽  
Author(s):  
Earl L. Smith ◽  
Yuzo Chino ◽  
Jinren Ni ◽  
Han Cheng
Keyword(s):  
Cortical Neurons ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Spatial Summation ◽  
Simple Cells ◽  
Complex Cells ◽  
Threshold Response ◽  
Binocular Interactions ◽  
Specific Complex ◽  
Left And Right

Smith, Earl L., III, Yuzo Chino, Jinren Ni, and Han Cheng. Binocular combination of contrast signals by striate cortical neurons in the monkey. J. Neurophysiol. 78: 366–382, 1997. With the use of microelectrode recording techniques, we investigated how the contrast signals from the two eyes are combined in individual cortical neurons in the striate cortex of anesthetized and paralyzed macaque monkeys. For a given neuron, the optimal spatial frequency, orientation, and direction of drift for sine wave grating stimuli were determined for each eye. The cell's disparity tuning characteristics were determined by measuring responses as a function of the relative interocular spatial phase of dichoptic stimuli that consisted of the optimal monocular gratings. Binocular contrast summation was then investigated by measuring contrast response functions for optimal dichoptic grating pairs that had left- to right-eye interocular contrast ratios that varied from 0.1 to 10. The goal was to determine the left- and right-eye contrast components required to produce a criterion threshold response. For all functional classes of cortical neurons and for both cooperative and antagonistic binocular interactions, there was a linear relationship between the left- and right-eye contrast components required to produce a threshold response. Thus, for example for cooperative binocular interactions, a reduction in contrast to one eye was counterbalanced by an equivalent increase in contrast to the other eye. These results showed that in simple cells and phase-specific complex cells, the contrast signals from the two eyes were linearly combined at the subunit level before nonlinear rectification. In non-phase-specific complex cells, the linear binocular convergence of contrast signals could have taken place either before or after the rectification process, but before spike generation. In addition, for simple cells, vector analysis of spatial summation showed that the inputs from the two eyes were also combined in a linear manner before nonlinear spike-generating mechanisms. Thus simple cells showed linear spatial summation not only within and between subregions in a given receptive field, but also between the left- and right-eye receptive fields. Overall, the results show that the effectiveness of a stimulus in producing a response reflects interocular differences in the relative balance of inputs to a given cell, however, the eye of origin of a light-evoked signal has no specific consequence.

Selective effects of aging on simple and complex cells in primary visual cortex of rhesus monkeys

2012 ◽  
Vol 1470 ◽  
pp. 17-23 ◽  
Author(s):  
Zhen Liang ◽  
Hongxin Li ◽  
Yun Yang ◽  
Guangxing Li ◽  
Yong Tang ◽  
...  
Keyword(s):  
Visual Cortex ◽  
Primary Visual Cortex ◽  
Rhesus Monkeys ◽  
Complex Cells ◽  
Simple And Complex Cells ◽  
Effects Of Aging ◽  
Selective Effects

Representing Response Position Relative to Display Location: Influence on Orthogonal Stimulus–Response Compatibility

2005 ◽  
Vol 58 (5) ◽  
pp. 839-864 ◽  
Author(s):  
Yang Seok Cho ◽  
Robert W. Proctor
Keyword(s):  
Irrelevant Stimulus ◽  
Stimulus Onset ◽  
The Body ◽  
Response Compatibility ◽  
Response Position ◽  
Stimulus Response ◽  
Stimulus Set ◽  
Response Sets ◽  
Stimulus Response Compatibility ◽  
Display Location

Two types of stimulus–response compatibility (SRC) effect occur with orthogonal stimulus and response sets, an overall up–right/down–left advantage and mapping preferences that vary with response position. Researchers agree that the former type is due to asymmetric coding of the stimulus and response alternatives, but disagree as to whether the latter type requires a different explanation in terms of the properties of the motor system. This issue is examined in three experiments. The location of the stimulus set influenced orthogonal SRC when it varied along the same dimension as the responses (Experiments 1 and 2), with the pattern predicted by the hypothesis that the stimulus set provides a referent relative to which response position is coded. The effect of stimulus-set location on orthogonal SRC was independent of the stimulus onset asynchrony (SOA) for a marker that indicated stimulus-set side and the imperative stimulus. In contrast, a spatial correspondence effect for the irrelevant stimulus-set location and response was a decreasing function of SOA. Experiment 3 showed that the orthogonal SRC effect was determined by response position relative to the stimulus-set location and not the body midline. The results support the view that both types of orthogonal SRC effects are due to asymmetric coding of the stimuli and responses.

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