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J W McClurkin

Publications and source records attributed to J W McClurkin.

At least 19 recordsLinked to original sources
PubMed1996

The influence of the visual cortex on the spatiotemporal response properties of lateral geniculate nucleus cells.

Previous studies of the cortical input to the mammalian dorsal lateral geniculate nucleus (LGN) have identified a number of possible functions for the corticogeniculate pathway, including alteration of LGN spatial frequency selectivity and facilitation of both binocular interactions and orientation selectivity. These changes may be due to either a tonic or a phasic cortical facilitation or both. The temporal differences between each of these inputs suggests that their impact on LGN cell temporal tuning should be unique. To test this hypothesis, we reversibly blocked the visual cortex (VI) and measured the effects on several indices of the temporal properties of LGN cells, including peak frequency, bandwidth, and response phase. Macaque monkeys were anesthetized and paralyzed during single cell recording from the LGN while area VI was cryogenically deactivated. Single-cell responses were visually evoked with drifting, luminance-modulated, sine-wave gratings and discrete-Fourier analyzed. Cortical cooling produced statistically significant increases or decreases in response amplitude in 64% of cells recorded. In most cases, alterations in response amplitude occurred for stimuli that varied in spatial as well as temporal frequency. For those cells influenced by changes in stimulus temporal frequency, the majority showed changes over a broad range of frequencies. A minority of cells showed changes in either peak temporal tuning or temporal frequency bandwidth. Response phase angles for all temporal frequencies tested were unaffected by cortical cooling. Overall, these results suggest that the cortical input may alter the temporal response properties of LGN cells, perhaps by tonic, but not exclusively excitatory, corticofugal influences.

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PubMed1996

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.

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PubMed1996

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.

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PubMed1991

Concurrent processing and complexity of temporally encoded neuronal messages in visual perception.

The intrinsic neuronal code that carries visual information and the perceptual mechanism for decoding that information are not known. However, multivariate statistics and information theory show that neurons in four visual areas simultaneously carry multiple, stimulus-related messages by utilizing multiplexed temporal codes. The complexity of these temporal messages increases progressively across the visual system, yet the temporal codes overlap in time. Thus, visual perception may depend on the concurrent processing of multiplexed temporal messages from all visual areas.

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PubMed1991

Visual responses of pulvinar and collicular neurons during eye movements of awake, trained macaques.

1. We recorded from single neurons in awake, trained rhesus monkeys in a lighted environment and compared responses to stimulus movement during periods of fixation with those to motion caused by saccadic or pursuit eye movements. Neurons in the inferior pulvinar (PI), lateral pulvinar (PL), and superior colliculus were tested. 2. Cells in PI and PL respond to stimulus movement over a wide range of speeds. Some of these cells do not respond to comparable stimulus motion, or discharge only weakly, when it is generated by saccadic or pursuit eye movements. Other neurons respond equivalently to both types of motion. Cells in the superficial layers of the superior colliculus have similar properties to those in PI and PL. 3. When tested in the dark to reduce visual stimulation from the background, cells in PI and PL still do not respond to motion generated by eye movements. Some of these cells have a suppression of activity after saccadic eye movements made in total darkness. These data suggest that an extraretinal signal suppresses responses to visual stimuli during eye movements. 4. The suppression of responses to stimuli during eye movements is not an absolute effect. Images brighter than 2.0 log units above background illumination evoke responses from cells in PI and PL. The suppression appears stronger in the superior colliculus than in PI and PL. 5. These experiments demonstrate that many cells in PI and PL have a suppression of their responses to stimuli that cross their receptive fields during eye movements. These cells are probably suppressed by an extraretinal signal. Comparable effects are present in the superficial layers of the superior colliculus. These properties in PI and PL may reflect the function of the ascending tectopulvinar system.

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PubMed1991

Lateral geniculate neurons in behaving primates. I. Responses to two-dimensional stimuli.

1. Using behaving monkeys, we studied the visual responses of single neurons in the parvocellular layers of the lateral geniculate nucleus (LGN) to a set of two-dimensional black and white patterns. We found that monkeys could be trained to make sufficiently reliable and stable fixations to enable us to plot and characterize the receptive fields of individual neurons. A qualitative examination of rasters and a statistical analysis of the data revealed that the responses of neurons were related to the stimuli. 2. The data from 5 of the 13 "X-like" neurons in our sample indicated the presence of antagonistic center and surround mechanisms and linear summation of luminance within center and surround mechanisms. We attribute the lack of evidence for surround antagonism in the eight neurons that failed to exhibit center-surround antagonism either to a mismatch between the size of the pixels in the stimuli and the size of the receptive field or to the lack of a surround mechanism (i.e., the type II neurons of Wiesel and Hubel). 3. The data from five other neurons confirm and extend previous reports indicating that the surround regions of X-like neurons can have nonlinearities. The responses of these neurons were not modulated when a contrast-reversing, bipartite stimulus was centered on the receptive field, which suggests a linear summation within the center and surround mechanisms. However, it was frequently the case for these neurons that stimuli of identical pattern but opposite contrast elicited responses of similar polarity, which indicates nonlinear behavior. 4. We found a wide variety of temporal patterns in the responses of individual LGN neurons, which included differences in the magnitude, width, and number of peaks of the initial on-transient and in the magnitude of the later sustained component. These different temporal patterns were repeatable and clearly different for different visual patterns. These results suggest that visual information may be carried in the shape as well as in the amplitude of the response waveform.

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PubMed1991

Lateral geniculate neurons in behaving primates. II. Encoding of visual information in the temporal shape of the response.

1. We used the Karhunen-Loève (K-L) transform to quantify the temporal distribution of spikes in the responses of lateral geniculate (LGN) neurons. The basis functions of the K-L transform are a set of waveforms called principal components, which are extracted from the data set. The coefficients of the principal components are uncorrelated with each other and can be used to quantify individual responses. The shapes of each of the first three principal components were very similar across neurons. 2. The coefficient of the first principal component was highly correlated with the spike count, but the other coefficients were not. Thus the coefficient of the first principal component reflects the strength of the response, whereas the coefficients of the other principal components reflect aspects of the temporal distribution of spikes in the response that are uncorrelated with the strength of the response. Statistical analysis revealed that the coefficients of up to 10 principal components were driven by the stimuli. Therefore stimuli govern the temporal distribution as well as the number of spikes in the response. 3. Through the application of information theory, we were able to compare the amount of stimulus-related information carried by LGN neurons when two codes were assumed: first, a univariate code based on response strength alone; and second, a multivariate temporal code based on the coefficients of the first three principal components. We found that LGN neurons were able to transmit an average of 1.5 times as much information using the three-component temporal code as they could using the strength code. 4. The stimulus set we used allowed us to calculate the amount of information each neuron could transmit about stimulus luminance, pattern, and contrast. All neurons transmitted the greatest amount of information about stimulus luminance, but they also transmitted significant amounts of information about stimulus pattern. This pattern information was not a reflection of the luminance or contrast of the pixel centered on the receptive field. 5. In addition to measuring the average amount of information each neuron transmitted about all stimuli, we also measured the amount of information each neuron transmitted about the individual stimuli with both the univariate spike count code and the multivariate temporal code. We then compared the amount of information transmitted per stimulus with the magnitudes of the responses to the individual stimuli. We found that the magnitudes of both the univariate and the multivariate responses to individual stimuli were poorly correlated with the information transmitted about the individual stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)

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PubMed1991

Lateral geniculate neurons in behaving primates. III. Response predictions of a channel model with multiple spatial-to-temporal filters.

1. For the experiments reported in these papers, we recorded the responses of lateral geniculate (LGN) neurons to a large set of two-dimensional, black and white patterns based on Walsh functions and to a set of test stimuli. In the first two papers we reported that these neurons encode stimulus-related information in both the strength and the shape of the response waveforms and that there are more than two independent components in the response. These results cannot be explained by existing models. This paper provides a model of LGN neurons that not only accounts for the foregoing observations, but also yields predictions confirmed by direct tests. 2. The model represents a neuron as a set of three parallel channels. The input to each channel is an array of pixel luminances. Each channel consists of an input nonlinearity cascaded into a linear spatial-to-temporal filter. The output of each channel is a basic waveform, a principal component. The response of the neuron is the sum of the outputs of the three channels. 3. The model accounted for much of the variance in the coefficients of the first three principal components of the neuronal responses to the set of Walsh stimuli. Using parameters derived from the responses of neurons to the Walsh stimuli only, the model also predicted the responses to "center-surround" annuli of different contrasts and mean luminances, as well as to superpositions of pairs of Walsh patterns. The model made statistically significant predictions of the coefficients of two of the principal components of these responses. 4. After the parameters of the model had been fit to reproduce the responses of neurons to the Walsh stimuli, we found that the input nonlinearity of the model was compressed at both the high and low luminance levels. This compression produced response saturation that closely resembled the response saturation of neurons reported in the first paper in this series. Although not absolutely smooth, the spatial filter for the first channel had a dominant excitatory or inhibitory center and an antagonistic surround. Thus this spatial filter accounted for both the center and the surround structures of previous models of LGN receptive fields. There was greater variety in the structures of the spatial filters for the second and third channels, but none had a center-surround organization. Many of the spatial filters for these higher channels contained oriented ridges or valleys. Other spatial filters were dominated by a bipolar pair of pixels. 5. The model of LGN neurons that we present in this paper represents an extension over previous models in four ways. First, the model is capable of explaining the responses of neurons to a wider range of luminances than previous models. Second, the model is capable of explaining the shapes of the response waveforms as well as their magnitudes. Third, the concept of a single receptive field is extended to a series of spatial-to-temporal filters. Fourth, the model suggests that LGN neurons provide a description of both the brightness and the form of a stimulus in their response waveforms.

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PubMed1990

Orbital position and eye movement influences on visual responses in the pulvinar nuclei of the behaving macaque.

We studied the influences of eye movements on the visual responses of neurons in two retinotopically organized areas of the pulvinar of the macaque. Cells were recorded from awake, trained monkeys, and visual responses were characterized immediately before and after the animals made saccadic eye movements. A significant proportion of the cells were more responsive to stimuli around the time of eye movements than they were at other intervals. Other cells had response reduction. For some neurons, the change in excitability was associated with orbital position and not the eye movement. For other cells the change was present with eye movements of similar amplitude and direction but with different starting and ending positions. Here it appears that the eye movement is the important parameter. Other cells had effects related to both eye position and eye movements. In all cells tested, the changes in excitability were present when the experiments were conducted in the dark as well as in the light. This suggests that the mechanism of the effect is related to the eye position or eye movement and not visual-visual interactions. For about half of the neurons with modulations, the response showed facilitation for stimuli presented in the most responsive region of the receptive field but not for those at the edge of the field. For the other cells there was facilitation throughout the field. Thus, a gradient of modulation in the receptive field may vary among cells. These experiments demonstrate modulations of visual responses in the pulvinar by eye movements. Such effects may be part of the visual-behavioral improvements at the end of eye movements and/or contribute to spatial constancy.

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PubMed1990

Abnormalities in visual spatial attention in men with mirror movements associated with isolated hypogonadotropic hypogonadism.

We studied spatial attentional performance on a visually cued reaction time task in men with isolated hypogonadotropic hypogonadism. A subset of these patients, who displayed mirror movements, have spatial attentional abnormalities. They were slow to respond to targets in the right visual field and especially slow when those targets followed incorrect or diffuse cues. This slowing was present for at least 500 msec after cue onset. They responded equally to targets in the left visual field independent of the spatial cues. The patient population as a whole was significantly faster than controls across all experimental conditions, although the speed of their attentional movement was normal. These data suggest that patients with isolated hypogonadotropic hypogonadism perform reaction time tasks quickly, that faster reaction times do not reflect superior attentional performance, and that a subset of these patients has a spatial attentional abnormality.

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PubMed1989

The visual superior colliculus and pulvinar.

The data reviewed here suggest several relationships between the superior colliculus, pulvinar and saccadic eye movements. These will be summarized in a series of steps in the visual initiation of saccadic eye movements. Area Pdm of the pulvinar appears to be involved in visual spatial attention. As such it may participate in the earliest decisions to make an eye movement: the shift of attention. This function is suggested by the spatially selective enhancement seen in the activity of individual neurons and the effects of drug injections on attentional performance. The attentional function of area Pdm is probably accomplished in coordination with cortical area 7 and possibly other areas. The spatially non-selective enhancement found in PI and PL may precede or follow the enhancement in Pdm. The next oculomotor function may be played by the visual cells in the superficial layers of the colliculus. Here the enhancement effect appears to be related to target selection which is specifically used with saccadic eye movements. Lesions here lead to animals who are not easily distracted by peripheral events; they do not select saccade targets. Thus the visual regions of the superior colliculus and Pdm may provide separate initiatives to the saccadic system. Once the eye begins to move, there are many relations manifested in both brain areas. Many of the visual collicular cells and those in parts of the pulvinar are unresponsive to visual stimuli during eye movements. In the colliculus, this particular effect may function to minimize erroneous visual targets whereas in the pulvinar the process may be more important perceptually. Finally there is a population of cells in the pulvinar which discharge during and after saccadic eye movements. These cells may have a role in the termination of one attentional scan and the beginning of the next. The pathways through which these effects may be mediated are totally unclear. The visual signal from the superficial layers related to saccade initiation may impinge directly on the cells in the intermediate layers or may reach the oculomotor system through other pathways. The parts of the pulvinar which relate to visual spatial attention are interconnected with posterior parietal cortex and probably function in concert with that area. How those data get to the oculomotor system has yet to be determined. Since the pulvinar is extensively interconnected with the cortex, it is most likely through this route that eye movement information coming from the superior colliculus is brought to higher perceptual areas.(ABSTRACT TRUNCATED AT 400 WORDS)

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PubMed1987

Release of cortical catecholamines by visual stimulation requires activity in thalamocortical afferents of monkey and cat.

Catecholamine (CA) release was measured in vivo in the monkey and cat visual cortices electrochemically. Stereate-modified, graphite-paste electrodes were used to monitor changes in norepinephrine and dopamine release. Micromolar changes in CA concentration were obtained by stimulation of the eye with nonspecific (strobe) or specific (oriented bars, radial gratings) stimuli. CA release depended on which eye was illuminated. Electrodes passed tangentially through the striate area recorded release following visual stimulation of one eye or the other in succession, and the shift in eye dominance occurred at about 500 microns intervals. The magnitude of CA release was highly correlated with the ocular dominance of neuronal activity measured with tungsten microelectrodes. Light-stimulated release was not recorded in monkey area V2, V4, or somatosensory area 1, but was recorded in cat V2, suggesting that the presence of LGN afferents is associated with CA release. Results are discussed in terms of the role of geniculate activity and the specific role of CAs in cortical information processing.

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PubMed1985

Evidence for spatial structure in the cortical input to the monkey lateral geniculate nucleus.

We combined visual stimulation of the corticogeniculate pathway with cryogenic blockade of area 17 to examine the visual spatial structure of the cortical influence on single macaque LGN cells. Excitatory central areas surrounded by inhibitory regions and vice versa were found with equal frequency. Unstructured influences were also seen. The structure of the influence suggests that modulation of the spatial parameter may be an important function for the corticogeniculate pathway.

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PubMed1984

Visual cortical input alters spatial tuning in monkey lateral geniculate nucleus cells.

The response of monkey lateral geniculate nucleus (l.g.n.) cells to flashing spots, annuli, and drifting sine-wave gratings were recorded with tungsten micro-electrodes. These stimuli were presented (a) monocularly, through an aperture in the centre of a radial grating, or (b) dichoptically, in which the spots or drifting gratings were presented to the dominant eye's receptive field, while the centre of the radial grating was positioned on the corresponding retinal location of the other eye. Movement of the radial grating produced changes in the l.g.n. cell responses evoked by the spots and sine-wave gratings. These changes were reversed by cryogenic blockade of the striate cortex. Therefore, radial grating movement altered the responses of l.g.n. cells by activating the corticogeniculate (c.g.) pathway. In about half of all cells, radial grating-induced alterations of centre, or surround, or both responses to spots and annuli were produced. By adopting a simple spatial filtering model of the centre and surround mechanisms, it was possible to predict how these alterations in centre/surround balance would affect the cell's responses to sine-wave gratings. Alterations were observed in the peak and band width of the spatial and/or temporal tuning curves. The radial gratings did not alter the spatial summation properties of cells. Minor alterations in the spectral neutral points of chromatically opponent neurones were occasionally found. These results are interpreted as support for the view that spatial and temporal tuning are dynamic properties of some l.g.n. neurones by virtue of descending input from the visual cortex.

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PubMed1982

Modulation of lateral geniculate nucleus cell responsiveness by visual activation of the corticogeniculate pathway.

A radial grating stimulus was used to assess the effect of stimulation of the region beyond the classical surround of monkey lateral geniculate nucleus (LGN) receptive fields. The effect was measured by the differences in the responsiveness of the LGN cell center to small flashing spots between two conditions: (1) grating stationary or (2) grating rotating. The grating was present only in regions beyond the classical center and surround. The rotating grating produced changes in the flash-evoked spike response but not in the spontaneous activity in about half of the X cells and all of the Y cells. The direction of the effect was independent of the sign of the receptive field center. In a control experiment, cryogenic blockade of striate cortex reversed the effect in all cells tested. The grating effect was still present for cells having fields in that part of visual space beyond the region represented by the cooled cortical area. The effect was not a result of activation of classical extra-receptive field influences, since cells showing the effect did not exhibit shift or periphery effects or outer disinhibitory surrounds. The effect was not seen in recordings from intrageniculate retinal axons. We conclude that the radial grating effects LGN cell responsivity by activation of the corticogeniculate pathway.

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PubMed1982

Spatial summation and conduction latency classification of cells of the lateral geniculate nucleus of macaques.

Cells in the lateral geniculate nucleus (LGN) of the macaque monkey were investigated with microelectrodes in an attempt to develop an overall classification scheme. We classified cells in the parvocellular (P) and magnocellular (M) layers according to (non)linearity of spatial summation, shock latency, and chromatic organization of center and surround. We also measured the spatial and temporal tuning to counterphasing and drifting sine wave gratings and tested for periphery effects. Our results showed that no strict laminar segregation existed for any cell property studied. Our results can be summarized as follows: 1. Most P layer cells showed a linear summation (98%) and color-opponent responses (80%), while other cells showed a nonlinear summation (Y-cells, 2%) and broad band responses (28%). In contrast, 37% of the M layer cells were linear summators and the remainder were nonlinear. Therefore, there are overlapping distributions of X- and Y-cells in P and M layers but not a strict segregation. 2. P layer cells had longer shock latencies than M layer cells. X-cells conducted more slowly (2.4 +/- 0.7 msec) than Y-cells (1.6 +/- 0.8 msec), but there were overlapping distributions. Latency shortened gradually, rather than abruptly, with increasing depth. 3. The first harmonic of X- and Y-cell responses was maximally sensitive to spatial frequencies of about 2 cycles/deg. Each type of cell modulated about a mean rate to a drifting grating, although Y-cells had higher distortion than X-cells. Response amplitudes to drifting gratings were higher for MX- and MY- than for PX-cells. No DC elevation to high spatial frequencies was seen. Spatial bandwidths averaged 2 to 5 octaves. X-cells were maximally tuned to temporal frequencies around 11 Hz, and Y-cells were tuned to about 19 Hz;. temporal bandwidths for both averaged 2.8 octaves. 4. Periphery effects were detected in 4% of the X-cells and 25% of the Y-cells. 5. These data indicate that gradual changes occur between dorsal and ventral layers: summation changes from linear to nonlinear; conduction latencies shorten; peak temporal tuning increases; response amplitudes increase; the periphery effect becomes more prevalent. Spatial tuning does not change. No strict laminar segregation or specificity exists for any of the properties that we studied.

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