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E Zohary

Publications and source records attributed to E Zohary.

12 recordsLinked to original sources

Correlated firing in macaque visual area MT: time scales and relationship to behavior.

We studied the simultaneous activity of pairs of neurons recorded with a single electrode in visual cortical area MT while monkeys performed a direction discrimination task. Previously, we reported the strength of interneuronal correlation of spike count on the time scale of the behavioral epoch (2 sec) and noted its potential impact on signal pooling (Zohary et al., 1994). We have now examined correlation at longer and shorter time scales and found that pair-wise cross-correlation was predominantly short term (10-100 msec). Narrow, central peaks in the spike train cross-correlograms were largely responsible for correlated spike counts on the time scale of the behavioral epoch. Longer-term (many seconds to minutes) changes in the responsiveness of single neurons were observed in auto-correlations; however, these slow changes in time were on average uncorrelated between neurons. Knowledge of the limited time scale of correlation allowed the derivation of a more efficient metric for spike count correlation based on spike timing information, and it also revealed a potential relative advantage of larger neuronal pools for shorter integration times. Finally, correlation did not depend on the presence of the visual stimulus or the behavioral choice of the animal. It varied little with stimulus condition but was stronger between neurons with similar direction tuning curves. Taken together, our results strengthen the view that common input, common stimulus selectivity, and common noise are tightly linked in functioning cortical circuits.

Action Potentials↗

Visuo-haptic object-related activation in the ventral visual pathway.

The ventral pathway is involved in primate visual object recognition. In humans, a central stage in this pathway is an occipito-temporal region termed the lateral occipital complex (LOC), which is preferentially activated by visual objects compared to scrambled images or textures. However, objects have characteristic attributes (such as three-dimensional shape) that can be perceived both visually and haptically. Therefore, object-related brain areas may hold a representation of objects in both modalities. Using fMRI to map object-related brain regions, we found robust and consistent somatosensory activation in the occipito-temporal cortex. This region showed clear preference for objects compared to textures in both modalities. Most somatosensory object-selective voxels overlapped a part of the visual object-related region LOC. Thus, we suggest that neuronal populations in the occipito-temporal cortex may constitute a multimodal object-related network.

Adult↗

Macaque monkeys categorize images by their ordinal number.

The recall of a list of items in a serial order is a basic cognitive skill. However, it is unknown whether a list of arbitrary items is remembered by associations between sequential items or by associations between each item and its ordinal position. Here, to study the nonverbal strategies used for such memory tasks, we trained three macaque monkeys on a delayed sequence recall task. Thirty abstract images, divided into ten triplets, were presented repeatedly in fixed temporal order. On each trial the monkeys viewed three sequentially presented sample stimuli, followed by a test stimulus consisting of the same three images and a distractor image (chosen randomly from the remaining 27). The task was to touch the three images in their original order without touching the distractor. The most common error was touching the distractor when it had the same ordinal number (in its own triplet) as the correct image. Thus, the monkeys' natural tendency was to categorize images by their ordinal number. Additional, secondary strategies were used eventually to avoid the distractor images. These included memory of the sample images (working memory) and associations between sequence triplet members. Thus, monkeys use multiple mnemonic strategies according to their innate tendencies and the requirements of the task.

Animals↗

Temporal gating of neural signals during performance of a visual discrimination task.

The flow of neural signals within the cerebral cortex must be subject to multiple controls as behaviour unfolds in time. In a visual discrimination task that includes a delay period, the transmission of sensory signals to circuitry that mediates memory, decision-making and motor-planning must be governed closely by 'filtering' or 'gating' mechanisms so that extraneous events occurring before, during or after presentation of the critical visual stimulus have little or no effect on the subject's behavioural responses. Here we study one such mechanism physiologically by applying electrical microstimulation to columns of directionally selective neurons in the middle temporal visual area at varying times during single trials of a direction-discrimination task. The behavioural effects of microstimulation varied strikingly according to the timing of delivery within the trial, indicating that signals produced by microstimulation may be subject to active 'gating'. Our results show several important features of this gating process: first, signal flow is modulated upwards on onset of the visual stimulus and downwards, typically with a slower time course, after stimulus offset; second, gating efficacy can be modified by behavioural training; and third, gating is implemented primarily downstream of the middle temporal visual area.

Animals↗

Inter-trial neuronal activity in inferior temporal cortex: a putative vehicle to generate long-term visual associations.

When monkeys perform a delayed match-to-sample task, some neurons in the anterior inferotemporal cortex show sustained activity following the presentation of specific visual stimuli, typically only those that are shown repeatedly. When sample stimuli are shown in a fixed temporal order, the few images that evoke delay activity in a given neuron are often neighboring stimuli in the sequence, suggesting that this delay activity may be the neural correlate of associative long-term memory. Here we report that stimulus-selective sustained activity is also evident following the presentation of the test stimulus in the same task. We use a neural network model to demonstrate that persistent stimulus-selective activity across the intertrial interval can lead to similar mnemonic representations (distributions of delay activity across the neural population) for neighboring visual stimuli. Thus, inferotemporal cortex may contain neural machinery for generating long-term stimulus-stimulus associations.

Animals↗

Integration across directions in dynamic random dot displays: vector summation or winner take all?

Recent studies have clearly demonstrated that the activity of directionally selective neuronal populations in the middle temporal (MT) and medial superior temporal (MST) cortical areas plays a direct role in the judgment of the direction of visual motion. However, the way in which the information is derived from a population of neurons remains unknown. Two principal models have been suggested in the past: the vector summation model suggests that the responses of neurons encoding all directions of motion are weighted and pooled to obtained an accurate estimate of the mean direction of motion; the winner-take-all model is based on a competition between different direction-specific channels, so that decisions are cast in favor of the channel generating the strongest directional signal. To discriminate between these two models we generated random dot stimuli that contained an asymmetric distribution of directions of motion. Human subjects were asked to adjust the global direction of motion to the upward vertical direction. When the directional signals were of similar strength, subjects tended to perceive global motion in the mean direction of motion (corresponding to vector summation), but as one directional signal became more prominent, most subjects' settings diverged from the mean towards the modal direction of motion. Some subjects could either match the mean or the modal direction of motion in the display, depending on the task instructions. These results suggest that the perceptual judgment of direction of motion is not based on any rigid algorithm generating a single valued output. Rather, human observers are able to judge different aspects of the distribution of activity in a cortical area depending on the task requirements.

Humans↗

Correlated neuronal discharge rate and its implications for psychophysical performance.

Single neurons can signal subtle changes in the sensory environment with surprising fidelity, often matching the perceptual sensitivity of trained psychophysical observers. This similarity poses an intriguing puzzle: why is psychophysical sensitivity not greater than that of single neurons? Pooling responses across neurons should average out noise in the activity of single cells, leading to substantially improved psychophysical performance. If, however, noise is correlated among these neurons, the beneficial effects of pooling would be diminished. To assess correlation within a pool, the responses of pairs of neurons were recorded simultaneously during repeated stimulus presentations. We report here that the observed covariation in spike count was relatively weak, the correlation coefficient averaging 0.12. A theoretical analysis revealed, however, that weak correlation can limit substantially the signalling capacity of the pool. In addition, theory suggests a relationship between neuronal responses and psychophysical decisions which may prove useful for identifying cell populations underlying specific perceptual capacities.

Action Potentials↗

Neuronal plasticity that underlies improvement in perceptual performance.

The electrophysiological properties of sensory neurons in the adult cortex are not immutable but can change in response to alterations of sensory input caused by manipulation of afferent pathways in the nervous system or by manipulation of the sensory environment. Such plasticity creates great potential for flexible processing of sensory information, but the actual effects of neuronal plasticity on perceptual performance are poorly understood. The link between neuronal plasticity and performance was explored here by recording the responses of directionally selective neurons in the visual cortex while rhesus monkeys practiced a familiar task involving discrimination of motion direction. Each animal experienced a short-term improvement in perceptual sensitivity during daily experiments; sensitivity increased by an average of 19 percent over a few hundred trials. The increase in perceptual sensitivity was accompanied by a short-term improvement in neuronal sensitivity that mirrored the perceptual effect both in magnitude and in time course, which suggests that improved psychophysical performance can result directly from increased neuronal sensitivity within a sensory pathway.

Animals↗

Mechanisms of velocity constancy.

Human observers can compare the physical velocities of objects (cm/sec) moving at different distances quite well, although the objects' retinal velocities (deg/sec) may vary considerably. This perceptual ability is called velocity constancy. We conducted a number of experiments to investigate what mechanisms observers use to attain this constancy and if pure motion signals can also be matched according to their physical speeds. Subjects were asked to match the velocities of two moving stimuli presented at different viewing distances. The stimuli consisted of sparse random-dot kinematograms or drifting Julesz patterns. The subjects matched the true physical velocities of the stimuli provided that the two visual scenes contained identical size references. Knowledge of the actual viewing distances proved to be irrelevant for evaluating the physical velocities of the stimuli. We conclude that velocity constancy is based upon a relative scaling algorithm.

Algorithms↗

Population coding of visual stimuli by cortical neurons tuned to more than one dimension.

Neurons in the visual cortex are typically selective to a number of stimulus dimensions. Thus, there is a basic ambiguity in relating the response level of a single neuron to the stimulus values. It is shown that a multi-dimensional stimulus may be coded reliably by an ensemble of neurons, using a weighted average population coding model. Each neurons' contribution to the population signal for each dimension is the product of its response magnitude and its preferred value for that dimension. The sum of the products was normalized by the sum of the ensemble responses. Simulation results show that the representation accuracy increases as the square root of the number of units irrespective of the number of dimensions. Comparison of a specific 2D case of this population code for orientation and spatial frequency to behavioral discrimination levels yields that 10(3)-10(4) neurons are needed to reach psychophysical performance. Introduction of each additional dimension requires about 1.7 times the number of neurons in the ensemble to reach the same level of accuracy. This result suggests that neurons may be selective for only 3 to 5 dimensions. It also provides another rationale for the existence of parallel processing streams in vision.

Animals↗

Time course of perceptual discrimination and single neuron reliability.

The reliability of identification of a visual target increases with time available for inspection of the stimulus. We suggest that the neural basis of this improvement is the existence of a mechanism for integrating a noisy firing rate over some period, leading to a reduction in mean firing rate variance with available processing time. We have determined the experimental time course of the improvement in reliability in a parallel search task where the available inspection time is limited by the presentation of a mask at various times after a brief stimulus. We compare the resulting psychometric functions with the predictions of a model based on Signal Detection Theory. The model is based on the assumption that the reliability of the observer's response is limited by the variability of the responses of individual neurons. The reliability of the discrimination between two stimuli at the neuronal level is then directly related to the ratio of the difference between their integrated mean responses (over many trials) to the response standard deviation. This reliability increases with inspection time. To demonstrate application of the model to electrophysiological data, "neurometric functions" are derived from the firing rates of a monkey V1 cortical neuron. The data were obtained while the animal was active in a discrimination task. The results correspond qualitatively to our observed human psychometric functions.

Animals↗

How serial is serial processing in vision?

Visual search for an element defined by the conjunction of its colour and orientation has previously been shown to be a serial processing task since reaction times increase linearly with the number of distractor elements used in the display. Evidence is presented that there are parallel processing constituents to this serial search. Processing time depended on the ratio of the number of the two distractor types used, suggesting that only one type was scanned. Which type was scanned also depended on the distractor ratio, indicating that this decision was made after stimulus presentation and was based on a parallel figure-ground separation of the stimulus elements. Furthermore, in accordance with this serial scanning model, there was an increase in processing speed (elements scanned per second) with increase in number of elements to be scanned. This increased efficiency suggests that clumps of elements were processed synchronously. Under the stimulation conditions used, clumps contained six to sixteen elements and each clump was processed in 50-150 ms.

Attention↗