Responses Of V1 Neurons Research Articles

Primate striate and prestriate cortical neurons during discrimination. I. simultaneous temporal encoding of information about color and pattern.

1. We recorded the responses of neurons in cortical areas V1, V2, and V4 to a set of 36 colored patterns while monkeys discriminated among the stimuli on the basis of their color or their pattern. In the discrimination task a colored square or a black and white pattern was presented foveally as a cue stimulus. The monkey was required to choose, by making a saccade, which of three peripheral targets had the same property as the cue. One of the peripheral targets was centered on the receptive field of the neuron, and the other two were positioned at equally distant points around the circumference of an imaginary circle centered on the cue and passing through the receptive field. 2. An examination of the responses to the stimuli showed that there was a complex interaction between the effects of color and of pattern on the neuronal responses. Because of these interactions, we tested sensitivity to color and pattern by sorting the responses to all stimuli according to the color or pattern of the stimulus. We found that the number of spikes in the responses was affected by only one or the other of the stimulus parameters, but that the temporal distribution of spikes was affected by both stimulus parameters. We quantified the relative sensitivities of each neuron to color and pattern by dividing the amount of information the neuron transmitted about color by the amount of information the neuron transmitted about pattern. The distributions of information ratios assuming a spike count code were broad, indicating that many neurons were sensitive to only one stimulus parameter or the other. In contrast, the distributions of information ratios assuming a wave-form code were narrow and centered near 1.0, indicating nearly equal sensitivities to both stimulus parameters. 3. In our initial experiments, it appeared that the color or pattern used as the cue for the discrimination task affected the responses of many neurons to stimuli on the receptive field. To determine whether the cue effect was due to simple visual interactions or to the cognitive requirements of the discrimination task, we performed a control experiment in which the cue was turned on 80 ms after the peripheral stimuli. For many of the neurons in the control experiment, an effect related to the cue appeared in the response before the cue had been turned on. Thus the effect we observed must have been due to visual interactions with the distractor targets, even though these were outside the neuron's classically defined receptive field. 4. We compared the rate at which color and pattern information developed in the response over time assuming either a spike count or a waveform code. The spike count code gained more of its information in the first 20ms of the response than did the waveform code, but thereafter the information carried by the spike count code developed more slowly and reached a lower asymptote than did the information carried by the waveform code. 5. The waveform codes carried nearly equal amounts of information about color and pattern, but the messages about these two parameters did not develop at the same rate in all areas. The messages about color and pattern developed at the same rate in area V1, but messages about color developed more slowly than did the messages about pattern in areas V2 and V4. 6. These results offer a neurophysiological basis for both the psychological separateness of color and pattern, and the binding of color and pattern into a unified percept. We propose that the separateness of color and form arises not by virtue of their being encoded by different populations of neurons, but by virtue of their being encoded by separable waveform codes in the responses of single neurons. We propose that the binding of color and form occurs by virtue of their codes being multiplexed on the same neurons.

Primate striate and prestriate cortical neurons during discrimination. II. separable temporal codes for color and pattern.

1. In the previous paper we reported our analysis of the responses of neurons in cortical areas V1, V2, and V4 to a set of stimuli that consisted of all 36 combinations of six colors and six patterns. Neurons in all three cortical areas simultaneously encoded information about both the color and pattern of the stimulus in the number and temporal distribution of spikes in their responses. To account for this ability, we propose that a neuron's response consists of separable temporal codes representing the color and pattern of the stimulus that are multiplexed together. 2. We used nonlinear regression to fit the model parameters to the data. We used the responses to 30 of the 36 stimuli as a training set to estimate the parameters of the model and the responses to the remaining 6 stimuli as a test set. After training, the model fitted the responses to stimuli in the training sets very well and predicted the responses to stimuli in the test sets. Thus neuronal responses to colored patterns contain separate temporal codes representing color and pattern. 3. After establishing the model parameters, we obtained the waveforms that represented each neuron's temporal codes for the six colors and six patterns of our stimulus set. We then proceeded with a series of analyses to determine whether these waveforms were viable candidates for neuronal codes. Cluster analysis revealed that there were only a few different classes of waveforms representing each color and pattern, and there were many neurons in each class. Further, neurons that used similar waveforms to represent one color or pattern also tended to use similar waveforms to represent other colors or patterns. The waveforms representing five of the six colors and three of the six patterns were similar in the two monkeys used in this study. 4. We compared the shapes of the code waveforms across cortical areas and found no differences among areas in the shapes of the waveforms representing four of the six colors. In contrast, we found that there were differences among areas in the shapes of the waveforms representing all six patterns. These results suggest that messages about color are encoded at an early level and are then propagated upward, but that messages about pattern are altered in each successive cortical area. 5. Our results offer a neurophysiological explanation for the psychophysical evidence that color and form are processed by different channels. We propose that the psychophysical channels for color and pattern arise from the separability of the temporal codes for color and pattern in the responses of single neurons. This hypothesis implies that psychophysical channels correspond to classes of temporal codes rather than to classes of neurons.

Focal attention produces spatially selective processing in visual cortical areas V1, V2, and V4 in the presence of competing stimuli.

1. The activity of single neurons was recorded in Macaca mulatta monkeys while they performed tasks requiring them to select a cued stimulus from an array of three to eight stimuli and report the orientation of that stimulus. Stimuli were presented in a circular array centered on the fixation target and scaled to place a single stimulus element within the receptive field of the neuron under study. The timing of the cuing event permitted the directing of visual attention to the spatial location of the correct stimulus before its presentation. 2. The effects of focal attention were examined in cortical visual areas V1, V2, and V4, where a total of 672 neurons were isolated with complete studies obtained for 94 V1, 74 V2, and 74 V4 neurons with receptive-field center eccentricities in the range 1.8-8 degrees. Under certain conditions, directed focal attention results in changes in the response of V1, V2, and V4 neurons to otherwise identical stimuli at spatially specific locations. 3. More than one-third of the neurons in each area displayed differential sensitivity when attention was directed toward versus away from the spatial location of the receptive field just before and during stimulus presentation. Both relative increases and decreases in neural activity were observed in association with attention directed at receptive-field stimuli. 4. The presence of multiple competing stimuli in the visual field was a major factor determining the presence or absence of differential sensitivity. About two-thirds of the neurons that were differentially sensitive to the attending condition in the presence of competing stimuli were not differentially sensitive when single stimuli were presented in control studies. For V1 and V2 neurons the presence of only a few (3-4) competing stimuli was sufficient for a majority of the neurons studied; a majority of the V4 neurons required six to eight stimuli in the array before significant differences between attending conditions occurred. 5. For V1 and V2 neurons the neuronal sensitivity differences between attending conditions were observed primarily at or near the peak of the orientation tuning sensitivity for each neuron; the differences were evident over a broader range of orientations in V4 neurons. 6. In conclusion, neural correlates of focal attentive processes can be observed in visual cortical processing in areas V1 and V2 as well as area V4 under conditions that require stimulus feature analysis and selective spatial processing within a field of competing stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)

Egocentric Spaw Representation in Early Vision

Abstract Recent physiological experiments have shown that the responses of many neurons in V1 and V3a are modulated by the direction of gaze. We have developed a neural network model of the hierarchy of maps in visual cortex to explore the hypothesis that visual features are encoded in egocentric (spatio-topic) coordinates at early stages of visual processing. Most psychophysical studies that have attempted to examine this question have concluded that features are represented in retinal coordinates, but the interpretation of these experiments does not preclude the type of retinospatiotopic representation that is embodied in our model. The model also explains why electrical stimulation experiments in visual cortex cannot distinguish between retinal and retinospatiotopic coordinates in the early stages of visual processing. Psychophysical predictions are made for testing the existence of retinospatiotopic representations.

Dynamics of responses of V1 neurons evoked by stimulation of different zones of receptive field

The dynamics of receptive fields of 73 neurons in area 17 of cat visual cortex were studied using the temporal slice method. Three-dimensional maps of the receptive fields were plotted using the criterion of spike number in successive fragments (step 10 or 20 ms) of responses to 100 local flashes presented at different parts of the receptive field in random order. The size and configuration of such dynamically recorded receptive fields were then estimated. This allowed us to reveal the dynamic reorganization of all receptive fields 20–400 ms after stimulation. A small zone of responses appeared in the receptive field after initial latency, then it widened, received definitive configuration, and after that decreased and disappeared. The effect was reproducible under repeated estimations. The relationships between receptive field and the previously described orientation tuning dynamics, as well as between dynamics of receptive fields and their summation zones, mechanisms and possible functional meaning of the revealed effects for signal processing in the primary visual cortex are discussed.

The response of area MT and V1 neurons to transparent motion

An important use of motion information is to segment a complex visual scene into surfaces and objects. Transparent motions present a particularly difficult problem for segmentation because more than one velocity vector occurs at each local region in the image, and current machine vision systems fail in these circumstances. The fact that motion transparency is prevalent in natural scenes, and yet artificial systems display an inability to analyze it, suggests that the primate visual system has developed specialized methods for perceiving transparent motion. Also, the currently prevalent model of physiological mechanisms for motion-direction selectivity employs inhibitory interactions between neurons; such interactions would silence neurons under transparent conditions and render the visual system blind to transparent motion. To examine how the primate visual system solves this transparency problem, we recorded the activity of direction-selective cells in the first (area V1) and in a later (area MT) stage in the cortical motion-processing pathway in behaving monkeys. The visual stimuli consisted of random dot patterns forming single moving surfaces, transparent surfaces, and motion discontinuities. We found that area V1 cells responded to their preferred direction of movement even under transparent conditions, whereas area MT cells were suppressed under the transparent condition. These data suggest a simple solution to the transparency problem at the level of area V1. More than one motion vector can be represented at a single retinal location by different subpopulations of neurons tuned to different directions of motion; these subpopulations may represent the early stage for segmenting different, transparent surfaces. The results also suggest that facilitatory mechanisms, which unlike inhibitory interactions are largely unaffected by transparent conditions, play an important role in direction selectivity in area V1. The inhibitory interactions for different motion directions for area MT neurons may contribute to a mechanism for smoothing or averaging the velocity field, computations thought to be necessary for reducing noise and interpolating moving surfaces from sparse information.

Pet birds as a risk factor for lung cancer.

The perceptual salience and visibility of image elements is influenced by other elements in their vicinity. The perceptual effect of image elements on an adjacent target element depends on their relative orientation. Collinear flanking elements usually improve sensitivity for the target element, whereas orthogonal elements have a weaker effect. It is believed that the collinear flankers exert these effects through lateral interactions between neurons in the primary visual cortex (area V1), but the precise mechanisms underlying these contextual interactions remain unknown. Here, we directly examined this question by recording the effects of flankers on the responses of V1 neurons at parafoveal representations while monkeys performed a fixation task or a contrast detection task. We found, unexpectedly, that collinear flankers reduce the monkeys9 perceptual sensitivity for a central target element. This behavioral effect was explained by a flanker-induced increase in the activity of V1 neurons in the absence of the central target stimulus, which reduced the amplitude of the target response. Our results indicate that the dominant effect of collinear flankers in parafoveal vision is suppression and suggest that these suppressive effects are caused by a decrease in the dynamic range of neurons coding the central target.

Contour, color and shape analysis beyond the striate cortex

The corticocortical pathway from striate cortex into the temporal lobe plays a crucial role in the visual recognition of objects. Anatomical studies indicate that this pathway is mainly organized as a serial hierarchy of multiple visual areas, including V1. V2. V3. V4. and inferior temporal cortex (IT). As expected from the anatomy, we have found that neurons in V4 and IT, like those in V1 and V2. are sensitive to many kinds of information relevant to object recognition. In the spatial domain, many V4 cells exhibit length, width, orientation, direction of motion and spatial frequency selectivity. In the spectral domain, many V4 cells are also tuned to wavelength. Thus, V4 is not specialized to analyze one particular attribute of a visual stimulus; rather, V4 appears to process both spatial and spectral information in parallel. A special contribution of V4 neurons to visual processing may lie in specific spatial and spectral interactions between their small excitatory receptive fields and large silent suppressive surrounds. Thus, although the excitatory receptive fields of V4 neurons are small, the responses of V4 neurons are influenced by stimuli throughout a much larger portion of the visual field. In IT. neurons also appear to process both spatial and spectral information throughout a large portion of the visual field. However, unlike V4 neurons, the excitatory receptive fields of IT neurons are very large. Many IT neurons, for example, are selective for the overall shape, color, or texture of a stimulus, anywhere within the central visual field. Together, these results suggest that within the areas of the occipito-temporal pathway, many different stimulus qualities are processed in parallel, but the type of analysis may become more global at each stage of processing.

RECEPTIVE FIELDS AND FUNCTIONAL ARCHITECTURE IN TWO NONSTRIATE VISUAL AREAS (18 AND 19) OF THE CAT.

RECEPTIVE FIELDS AND FUNCTIONAL ARCHITECTURE IN TWO NONSTRIATE VISUAL AREAS (18 AND 19) OF THE CAT.