The temporal resolution of neural codes: does response latency have a unique role?

This article reviews the nature of the neural code in non-human primate cortex and assesses the potential for neurons to carry two or more signals simultaneously. Neurophysiological recordings from visual and motor systems indicate that the evidence for a role for precisely timed spikes relative to other spike times (ca. 1-10 ms resolution) is inconclusive. This indicates that the visual system does not carry a signal that identifies whether the responses were elicited when the stimulus was attended or not. Simulations show that the absence of such a signal reduces, but does not eliminate, the increased discrimination between stimuli that are attended compared with when the stimuli are unattended. The increased accuracy asymptotes with increased gain control, indicating limited benefit from increasing attention. The absence of a signal identifying the attentional state under which stimuli were viewed can produce the greatest discrimination between attended and unattended stimuli. Furthermore, the greatest reduction in discrimination errors occurs for a limited range of gain control, again indicating that attention effects are limited. By contrast to precisely timed patterns of spikes where the timing is relative to other spikes, response latency provides a fine temporal resolution signal (ca. 10 ms resolution) that carries information that is unavailable from coarse temporal response measures. Changes in response latency and changes in response magnitude can give rise to different predictions for the patterns of reaction times. The predictions are verified, and it is shown that the standard method for distinguishing executive and slave processes is only valid if the representations of interest, as evidenced by the neural code, are known. Overall, the data indicate that the signalling evident in neural signals is restricted to the spike count and the precise times of spikes relative to stimulus onset (response latency). These coding issues have implications for our understanding of cognitive models of attention and the roles of executive and slave systems.

Excess synchrony in motor cortical neurons provides redundant direction information with that from coarse temporal measures.

Previous studies have shown that measures of fine temporal correlation, such as synchronous spikes, across responses of motor cortical neurons carries more directional information than that predicted from statistically independent neurons. It is also known, however, that the coarse temporal measures of responses, such as spike count, are not independent. We therefore examined whether the information carried by coincident firing was related to that of coarsely defined spike counts and their correlation. Synchronous spikes were counted in the responses from 94 pairs of simultaneously recorded neurons in primary motor cortex (MI) while monkeys performed arm movement tasks. Direct measurement of the movement-related information indicated that the coincident spikes (1- to 5-ms precision) carry approximately 10% of the information carried by a code of the two spike counts. Inclusion of the numbers of synchronous spikes did not add information to that available from the spike counts and their coarse temporal correlation. To assess the significance of the numbers of coincident spikes, we extended the stochastic spike count matched (SCM) model to include correlations between spike counts of the individual neural responses and slow temporal dependencies within neural responses (approximately 30 Hz bandwidth). The extended SCM model underestimated the numbers of synchronous spikes. Therefore as with previous studies, we found that there were more synchronous spikes in the neural data than could be accounted for by this stochastic model. However, the SCM model accounts for most (R(2) = 0.93 +/- 0.05, mean +/- SE) of the differences in the observed number of synchronous spikes to different directions of arm movement, indicating that synchronous spiking is directly related to spike counts and their broad correlation. Further, this model supports the information theoretic analysis that the synchronous spikes do not provide directional information beyond that available from the firing rates of the same pool of directionally tuned MI neurons. These results show that detection of precisely timed spike patterns above chance levels does not imply that those spike patterns carry information unavailable from coarser population codes but leaves open the possibility that excess synchrony carries other forms of information or serves other roles in cortical information processing not studied here.

Consistency of encoding in monkey visual cortex.

Are different kinds of stimuli (for example, different classes of geometric images or naturalistic images) encoded differently by visual cortex, or are the principles of encoding the same for all stimuli? We examine two response properties: (1) the range of spike counts that can be elicited from a neuron in epochs representative of short periods of fixation (up to 400 msec), and (2) the relation between mean and variance of spike counts elicited by different stimuli, that together characterize the information processing capabilities of a neuron using the spike count code. In monkey primary visual cortex (V1) complex cells, we examine responses elicited by static stimuli of four kinds (photographic images, bars, gratings, and Walsh patterns); in area TE of inferior temporal cortex, we examine responses elicited by static stimuli in the sample, nonmatch, and match phases of a delayed match-to-sample task. In each area, the ranges of mean spike counts and the relation between mean and variance of spike counts elicited are sufficiently similar across experimental conditions that information transmission is unaffected by the differences across stimulus set or behavioral conditions [although in 10 of 27 (37%) of the V1 neurons there are statistically significant but small differences, the median difference in transmitted information for these neurons was 0.9%]. Encoding therefore appears to be consistent across experimental conditions for neurons in both V1 and TE, and downstream neurons could decode all incoming signals using a single set of rules.

Effect of image orientation and size on object recognition: responses of single units in the macaque monkey temporal cortex.

This study examined how cells in the temporal cortex code orientation and size of a complex object. The study focused on cells selectively responsive to the sight of the head and body but unresponsive to control stimuli. The majority of cells tested (19/26, 73%) were selectively responsive to a particular orientation in the picture plane of the static whole body stimulus, 7/26 cells showed generalisation responding to all orientations (three cells with orientation tuning superimposed on a generalised response). Of all cells sensitive to orientation, the majority (15/22, 68%) were tuned to the upright image. The majority of cells tested (81%, 13/16) were selective for stimulus size. The remaining cells (3/16) showed generalisation across four-fold decrease in size from life-sized. All size-sensitive cells were tuned to life-sized stimuli with decreasing responses to stimuli reduced from life-size. These results do not support previous suggestions that cells responsive to the head and body are selective to view but generalise across orientation and size. Here, extensive selectivity for size and orientation is reported. It is suggested that object orientation and size-specific responses might be pooled to obtain cell responses that generalise across size and orientation. The results suggest that experience affects neuronal coding of objects in that cells become tuned to views, orientation, and image sizes that are commonly experienced. Models of object recognition are discussed.

I see a face-a happy face.

Information analysis shows that face-selective neurons in inferior temporal cortex encode different stimulus attributes early and late in their response to the same image.

Stochastic nature of precisely timed spike patterns in visual system neuronal responses.

It is not clear how information related to cognitive or psychological processes is carried by or represented in the responses of single neurons. One provocative proposal is that precisely timed spike patterns play a role in carrying such information. This would require that these spike patterns have the potential for carrying information that would not be available from other measures such as spike count or latency. We examined exactly timed (1-ms precision) triplets and quadruplets of spikes in the stimulus-elicited responses of lateral geniculate nucleus (LGN) and primary visual cortex (V1) neurons of the awake fixating rhesus monkey. Large numbers of these precisely timed spike patterns were found. Information theoretical analysis showed that the precisely timed spike patterns carried only information already available from spike count, suggesting that the number of precisely timed spike patterns was related to firing rate. We therefore examined statistical models relating precisely timed spike patterns to response strength. Previous statistical models use observed properties of neuronal responses such as the peristimulus time histogram, interspike interval, and/or spike count distributions to constrain the parameters of the model. We examined a new stochastic model, which unlike previous models included all three of these constraints and unlike previous models predicted the numbers and types of observed precisely timed spike patterns. This shows that the precise temporal structures of stimulus-elicited responses in LGN and V1 can occur by chance. We show that any deviation of the spike count distribution, no matter how small, from a Poisson distribution necessarily changes the number of precisely timed spike patterns expected in neural responses. Overall the results indicate that the fine temporal structure of responses can only be interpreted once all the coarse temporal statistics of neural responses have been taken into account.

Response features determining spike times.

Interpreting messages encoded in single neuronal responses requires knowing which features of the responses carry information. That the number of spikes is an important part of the code has long been obvious. In recent years, it has been shown that modulation of the firing rate with about 25 ms precision carries information that is not available from the total number of spikes across the whole response. It has been proposed that patterns of exactly timed (1 ms precision) spikes, such as repeating triplets or quadruplets, might carry information that is not available from knowing about spike count and rate modulation. A model using the spike count distribution, the low-pass filtered PSTH (bandwidth below 30 Hz), and, to a small degree, the interspike interval distribution predicts the numbers and types of exactly-timed triplets and quadruplets that are indistinguishable from those found in the data. From this it can be concluded that the coarse (< 30 Hz) sequential correlation structure over time gives rise to the exactly timed patterns present in the recorded spike trains. Because the coarse temporal structure predicts the fine temporal structure, the information carried by the fine temporal structure must be completely redundant with that carried by the coarse structure. Thus, the existence of precisely timed spike patterns carrying stimulus-related information does not imply control of spike timing at precise time scales.

Visual recognition based on temporal cortex cells: viewer-centred processing of pattern configuration.

A model of recognition is described based on cell properties in the ventral cortical stream of visual processing in the primate brain. At a critical intermediate stage in this system, 'Elaborate' feature sensitive cells respond selectively to visual features in a way that depends on size (+/- 1 octave), orientation (+/- 45 degrees) but does not depend on position within central vision (+/- 5 degrees). These features are simple conjunctions of 2-D elements (e.g. a horizontal dark area above a dark smoothly convex area). They can arise either as elements of an object's surface pattern or as a 3-D component bounded by an object's external contour. By requiring a combination of several such features without regard to their position within the central region of the visual image, 'Pattern' sensitive cells at higher levels can exhibit selectivity for complex configurations that typify objects seen under particular viewing conditions. Given that input features to such Pattern sensitive cells are specified in approximate size and orientation, initial cellular 'representations' of the visual appearance of object type (or object example) are also selective for orientation and size. At this level, sensitivity to object view (+/- 60 degrees) arises because visual features disappear as objects are rotated in perspective. Processing is thus viewer-centred and the neurones only respond to objects seen from particular viewing conditions or 'object instances'. Combined sensitivity to multiple features (conjunctions of elements) independent of their position, establishes selectivity for the configurations of object parts (from one view) because rearranged configurations of the same parts yield images lacking some of the 2-D visual features present in the normal configuration. Different neural populations appear to be selectively tuned to particular components of the same biological object (e.g. face, eyes, hands, legs), perhaps because the independent articulation of these components gives rise to correlated activity in different sets of input visual features. Generalisation over viewing conditions for a given object can be established by hierarchically pooling outputs of view-condition specific cells with pooling operations dependent on the continuity in experience across viewing conditions. Different object parts are seen together and different views are seen in succession when the observer walks around the object. The view specific coding that characterises the selectivity of cells in the temporal lobe can be seen as a natural consequence of selective experience of objects from particular vantage points. View specific coding for the face and body also has great utility in understanding complex social signals, a property that may not be feasible with object-centred processing.

Evidence accumulation in cell populations responsive to faces: an account of generalisation of recognition without mental transformations.

In this paper we analyse the time course of neuronal activity in temporal cortex to the sight of the head and body. Previous studies have already demonstrated the impact of view, orientation and part occlusion on individual cells. We consider the cells as a population providing evidence in the form of neuronal activity for perceptual decisions related to recognition. The time course on neural responses to stimuli provides an explanation of the variation in speed of recognition across different viewing circumstances that is seen in behavioural experiments. A simple unifying explanation of the behavioural effects is that the speed of recognition of an object depends on the rate of accumulation of activity from neurones selective for the object, evoked by a particular viewing circumstance. This in turn depends on the extent that the object has been seen previously under the particular circumstance. For any familiar object, more cells will be tuned to the configuration of the object's features present in the view or views most frequently experienced. Therefore, activity amongst the population of cells selective for the object's appearance will accumulate more slowly when the object is seen in an unusual view, orientation or size. This accounts for the increased time to recognise rotated views without the need to postulate 'mental rotation' or 'transformations' of novel views to align with neural representations of familiar views.

The 'Ideal Homunculus': decoding neural population signals.

Information processing in the nervous system involves the activity of large populations of neurons. It is possible, however, to interpret the activity of relatively small numbers of cells in terms of meaningful aspects of the environment. 'Bayesian inference' provides a systematic and effective method of combining information from multiple cells to accomplish this. It is not a model of a neural mechanism (neither are alternative methods, such as the population vector approach) but a tool for analysing neural signals. It does not require difficult assumptions about the nature of the dimensions underlying cell selectivity, about the distribution and tuning of cell responses or about the way in which information is transmitted and processed. It can be applied to any parameter of neural activity (for example, firing rate or temporal pattern). In this review, we demonstrate the power of Bayesian analysis using examples of visual responses of neurons in primary visual and temporal cortices. We show that interaction between correlation in mean responses to different stimuli (signal) and correlation in response variability within stimuli (noise) can lead to marked improvement of stimulus discrimination using population responses.

Gaze following and joint attention in rhesus monkeys (Macaca mulatta).

Gaze and attention direction provide important sources of social information for primates. Behavioral studies show that chimpanzees spontaneously follow human gaze direction. By contrast, non-ape species such as macaques fail to follow gaze cues. The authors investigated the reactions of rhesus macaques (Macaca mulatta) to attention cues of conspecifics. Two subjects were presented with videotaped images of a stimulus monkey with its attention directed to 1 of 2 identical objects. Analysis of eye movements revealed that both subjects inspected the target (object or position attended by the stimulus monkey) more often than the distractor (nonattended object or position). These results provide evidence that rhesus monkeys follow gaze and use the attention cues of other monkeys to orient their own attention to objects.

Integration of form and motion in the anterior superior temporal polysensory area (STPa) of the macaque monkey.

1. Processing of visual information in primates is believed to occur in at least two separate cortical pathways, commonly labeled the "form" and "motion" pathways. This division lies in marked contrast to our everyday visual experience, in which we have a unified percept of both the form and motion of objects, implying integration of both types of information. We report here on a neuronal population in the anterior part of the superior temporal polysensory area (STPa) both sensitive to form (heads and bodies) and selective for motion direction. 2. A total of 161 cells were found to be sensitive to body form and motion. The majority of cells (125 of 161, 78%) responded to only one combination of view and direction (termed unimodal cells, e.g., left profile view moving left, not right profile moving left, or left profile moving right). We show that the response of some of these cells is selective for both the motion and the form of a single object, not simply the juxtaposition of appropriate form and motion signals. 3. A smaller number of cells (9 of 161, 6%) responded selectively to two opposite combinations of view and direction (e.g., left profile moving left and right profile moving right, but no other view and direction combinations). A few cells (4 of 161, 2%) showed "object-centered" selectivity to view and direction combinations, responding to all directions of motion where the body moves in a direction compatible with the direction it faces, for example, responding to left profile going left, right profile going right, face view moving toward the observer, back view moving away from the observer, but not other view and direction combinations. 4. The majority of the neurons (106 of 138, 77%) selective for specific body view and direction combinations responded best to compatible motion (e.g., left profile moving left), and one fourth (23%) showed selectivity for incompatible motion (e.g., right profile moving left). 5. The relative strengths of motion and form inputs to cells in STPa conjointly sensitive to information about form and motion were assessed. The majority of the responses (95%) were characterized as showing nonlinear summation of form and motion inputs. 6. The capacity to discriminate different directions and different forms was compared across three populations of STPa cells, namely those sensitive to 1) form only, 2) motion only, and 3) both form and motion. The selectivity of the latter class could be predicted from combinations of the other two classes. 7. The response latencies of cells selective for form and motion are on average coincident with cells selective for direction of motion (but not stimulus form). Both these cell populations have response latencies on average 20 ms earlier than cells selective for static form. 8. Calculation of the average of early response latency cells (cell whose response latency was under the sample mean) suggests that direction information is present in cell responses some 35 ms before form information becomes evident. Direction information and form information become evident within 5 ms of each other in the average late response latency cells (those cells whose response latency was greater than the sample mean). Inputs relating to movement show an initial response period that does not discriminate direction. The quality of initial direction discrimination appeared to be independent of response latency. The initial discrimination of form was related to response latency in that cells with longer response latencies showed greater initial discrimination of form in their responses. We argue that these findings are consistent with form inputs arriving to area STPa approximately 20 ms after motion inputs into area STPa.

Recognition of objects and their component parts: responses of single units in the temporal cortex of the macaque.

We investigated the role that different component parts play in the neural encoding of the visual appearance of one complex object in the temporal cortex. Cells responsive to the sight of the entire human body (but no to control stimuli) were tested with two subregions (head alone with the body occluded from sight and the body alone with the head occluded). Forty-two percent (22 of 53) of cells responded to the whole body and to one of the two body regions tested separately: 72% (17 of 22) responding to the head and 28% (5 of 22) to the rest of the body. Forty-two percent (22 of 53) of cells responded independently to both regions of the body when tested in isolation. The remaining cells (17%, 9 of 53) were selective for the entire body and unresponsive to component parts. The majority of cells tested (90%, 35 of 39) were selective for perspective view (e.g., some cells respond optimally to the side view of the body, others to the back view). Comparable levels of view sensitivity were found for responses to the whole body and its parts. Results indicate (1) separate neuronal analysis of body parts and (2) extensive integration of information from different parts. Contrary to influential models of object recognition (Marr and Nishihara, 1978; Biederman, 1987), the results indicate view-specific processing both for the appearance of separate object components and for integration of information across components.

Responses of Anterior Superior Temporal Polysensory (STPa) Neurons to "Biological Motion" Stimuli.

Abstract Cells have been found in the superior temporal polysensory area (STPa) of the macaque temporal cortex that are selectively responsive to the sight of particular whole body movements (e.g., walking) under normal lighting. These cells typically discriminate the direction of walking and the view of the body (e.g., left profile walking left). We investigated the extent to which these cells are responsive under "biological motion" conditions where the form of the body is defined only by the movement of light patches attached to the points of limb articulation. One-third of the cells (25/72) selective for the form and motion of walking bodies showed sensitivity to the moving light displays. Seven of these cells showed only partial sensitivity to form from motion, in so far as the cells responded more to moving light displays than to moving controls but failed to discriminate body view. These seven cells exhibited directional selectivity. Eighteen cells showed statistical discrimination for both direction of movement and body view under biological motion conditions. Most of these cells showed reduced responses to the impoverished moving light stimuli compared to full light conditions. The 18 cells were thus sensitive to detailed form information (body view) from the pattern of articulating motion. Cellular processing of the global pattern of articulation was indicated by the observations that none of these cells were found sensitive to movement of individual limbs and that jumbling the pattern of moving limbs reduced response magnitude. A further 10 cells were tested for sensitivity to moving light displays of whole body actions other than walking. Of these cells 5/10 showed selectivity for form displayed by biological motion stimuli that paralleled the selectivity under normal lighting conditions. The cell responses thus provide direct evidence for neural mechanisms computing form from nonrigid motion. The selectivity of the cells was for body view, specific direction, and specific type of body motion presented by moving light displays and is not predicted by many current computational approaches to the extraction of form from motion.

Directional tuning of motion-sensitive cells in the anterior superior temporal polysensory area of the macaque.

An investigation was made into the directional sensitivity of cells in the macaque anterior superior temporal polysensory region (STPa) to the motion of objects. The cells studied were sensitive to the presence of motion but showed little or no selectivity for the form of the stimulus. Directional tuning was not continuously distributed about all possible directions. The majority of cells were most responsive to motion in a direction within 15 degrees of one of the three cartesian axes (up/down, left/right, towards/away). Tuning to direction varied in sharpness. For most (34/37) cells the angular change in direction required to reduce response to half maximal was between 45 and 70 degrees (for 3/37 cells it was > 90 degrees). The estimates of the directionality (median Id = 0.97) of STPa cells was similar to that reported for posterior motion processing areas (the middle temporal area, MT, and the medial superior temporal area, MST). The tuning for direction (sharpness, distribution and discrimination) of the motion-sensitive STPa cells were found to be similar to the tuning for perspective view of STPa cells selective for static form of the head and body. On average the STPa responses showed a 100- to 300-ms transient burst of activity followed by a tonic discharge maintained at approximately 20% of the peak firing rate for the duration of stimulation. The responses of motion-sensitive STPa cells occurred at an earlier latency (mean 91 ms) than responses of cells selective for static form (mean 119 ms), but the time course of responses of the two classes of cell were similar in many other respects. The early response latency and directional selectivity indicate that motion sensitivity in STPa cells derives from the dorsal visual pathway via MT/MST. The similarity of tuning for direction and perspective view within STPa may facilitate the integration of motion and form processing within this high-level brain area.

Time course of neural responses discriminating different views of the face and head.

1. Measurements of the magnitude and time course of response were made from 44 cells responsive to static head views at different levels of stimulus effectiveness. In this way responses to complex stimulus patterns evoking good, poor, and midrange responses could be compared across the cell population. 2. Cells exhibiting both good and poor initial discrimination between head views were found at short and long latencies; there was no correlation of any of the temporal response parameters measured with cell response latency. 3. The time course of the population response to the most effective stimuli showed a rapid increase to a peak firing rate (onset to peak, rise time, 58 ms) that was on average 115 spikes/s above spontaneous activity (S/A), followed by slower decay (decay time, 93 ms) to a maintained discharge rate (15% of the peak rate above S/A). 4. Discrimination between responses to different head views exhibited by the population showed a sharp rise and reached highly significant levels within 25 ms after the population's response onset. 5. On average, activity in a single neuron (the Average Cell) rises to 44% of its peak response rate within 5 ms of the response onset. 6. The Average Cell also showed exceptionally fast discrimination between views, significant within 5 ms of response onset. 7. It is argued that the fast rise in firing rate, followed by a decay to a lower rate and the very fast emergence of discrimination are features of pattern processing present in real neural systems that are lacking in many processing models based on artificial networks of neuronlike elements, particularly those where discrimination relies on top-down and/or lateral competitive inhibition. 8. It is concluded that the only way to account for the rapid discrimination is to consider a coding system in which the first spike from multiple sources is used to transmit information between stages of processing.

Organization and functions of cells responsive to faces in the temporal cortex.

Cells selectively responsive to the face have been found in several visual sub-areas of temporal cortex in the macaque brain. These include the lateral and ventral surfaces of inferior temporal cortex and the upper bank, lower bank and fundus of the superior temporal sulcus (STS). Cells in the different regions may contribute in different ways to the processing of the facial image. Within the upper bank of the STS different populations of cells are selective for different views of the face and head. These cells occur in functionally discrete patches (3-5 mm across) within the STS cortex. Studies of output connections from the STS also reveal a modular anatomical organization of repeating 3-5 mm patches connected to the parietal cortex, an area thought to be involved in spatial awareness and in the control of attention. The properties of some cells suggest a role in the discrimination of heads from other objects, and in the recognition of familiar individuals. The selectivity for view suggests that the neural operations underlying face or head recognition rely on parallel analyses of different characteristic views of the head, the outputs of these view-specific analyses being subsequently combined to support view-independent (object-centred) recognition. An alternative functional interpretation of the sensitivity to head view is that the cells enable an analysis of 'social attention', i.e. they signal where other individuals are directing their attention. A cell maximally responsive to the left profile thus provides a signal that the attention (of another individual) is directed to the observer's left. Such information is useful for analysing social interactions between other individuals.(ABSTRACT TRUNCATED AT 250 WORDS)

The effects of lighting conditions on responses of cells selective for face views in the macaque temporal cortex.

Neural mechanisms underlying recognition of objects must overcome the changes in an object's appearance caused by inconsistent viewing conditions, particularly those that occur with changes in lighting. In humans, lesions to the posterior visual association cortex can impair the ability to recognize objects and faces across different lighting conditions. Inferotemporal lesions in monkey have been shown to produce a similar difficulty in object matching tasks. Here we report on the extent to which cell responses selective for the face and other views of the head in monkey temporal cortex tolerate changes in lighting. For each cell studied the (preferred) head view eliciting maximal response was first established under normal lighting. Cells were then tested with the preferred head view lit from different directions (i.e. front, above, below or from the side). Responses of some cells failed to show complete generalization across all lighting conditions but together as a "population" they responded equally strongly under all four lighting conditions. Further tests on sub-groups of cells revealed that stimulus selectivity was maintained despite unusual lighting. The cells discriminated between head and control stimuli and between different views of the head independent of the lighting direction. The results indicate that constancy of recognition across different lighting conditions is apparent in the responses of single cells in the temporal cortex. Lighting constancy appears to be established by matching the retinal image to view-specific descriptions of objects (i.e. neurons which compute object structure from a limited range of perspective views).

Viewer-centred and object-centred coding of heads in the macaque temporal cortex.

An investigation was made into the sensitivity of cells in the macaque superior temporal sulcus (STS) to the sight of different perspective views of the head. This allowed assessment of (a) whether coding was 'viewer-centred' (view specific) or 'object-centred' (view invariant) and (b) whether viewer-centred cells were preferentially tuned to 'characteristic' views of the head. The majority of cells (110) were found to be viewer-centred and exhibited unimodal tuning to one view. 5 cells displayed object-centred coding responding equally to all views of the head. A further 5 cells showed 'mixed' properties, responding to all views of the head but also discriminating between views. 6 out of 56 viewer and object-centred cells exhibited selectivity for face identity or species. Tuning to view varied in sharpness. For most (54/73) cells the angle of perspective rotation reducing response to half maximal was 45-70 degrees but for 19/73 it was greater than 90 degrees. More cells were optimally tuned to characteristic views of the head (the full face or profile) than to other views. Some cells were, however, found tuned to intermediate views throughout the full 360 degree range. This coding of many distinct head views may have a role in the analysis of social signals based on the interpretation of the direction of other individuals' attention.