Biomedical subjects

E T Rolls

Publications and source records attributed to E T Rolls.

At least 127 records · Page 7Linked to original sources

PubMed1989

The effect of learning on the face selective responses of neurons in the cortex in the superior temporal sulcus of the monkey.

Neurophysiological studies have shown that some neurons in the cortex in the superior temporal sulcus and the inferior temporal gyrus of macaque monkeys respond to faces. These neurons provided a consistently identifiable substrate with which studies of the storage of visual information were performed. To determine whether face responsive neurons change how much they respond to different novel faces as they become familiar, neurons were tested with two experimental designs. In the first experiment, 22 neurons were tested on their responsiveness to the different members of a large set of novel faces as the set was presented repeatedly until the faces became familiar. 6 neurons altered the relative degree to which they responded to the different members of the set between the first two presentations and subsequent presentations. In a control condition, only 1 out of 17 neurons showed a significant response difference between the first two presentations and subsequent presentations when the experiment started with faces which were already familiar to the monkey. In the second experiment, 26 neurons were tested on their responsiveness to the different members of a set of familiar faces before and after the addition of a novel face to the set. 5 neurons altered the relative degree in which they responded to the different members of the set of familiar faces after addition of a novel face. It is suggested that these changes in neuronal responsiveness to different stimuli reflect the setting up of an ensemble encoded representation of face stimuli. This alteration of neuronal responsiveness as novel faces become familiar suggests that face responsive neurons may store information useful in visual recognition. In addition to this relatively long-term alteration of relative neuronal responsiveness to different stimuli, it was found that a large number of cells showed a higher mean response to the first presentation of a set of novel faces than to subsequent presentations of the faces. However, the response to the first presentation of a set of familiar faces was also higher than to subsequent presentations in that sequence. This pattern indicates a short term recency effect in the response of these neurons to visual stimuli which is similar to that previously reported (Baylis and Rolls 1987).

Animals↗

PubMed1989

Activity of hippocampal formation neurons in the monkey related to a conditional spatial response task.

To analyze neurophysiologically the functions of the primate hippocampus, the activity of 905 single hippocampal formation neurons was analyzed in two rhesus monkeys performing a conditional spatial response task known to be impaired in monkeys and in man by damage to the hippocampus or fornix. In the task, the monkey learned to make one spatial response, touching a screen three times when he saw one visual stimulus on the video monitor, and a different spatial response, of withdrawing his hand from the screen, when a different visual stimulus was shown. Fourteen percent of the neurons fired differentially to one or the other of the stimulus-spatial response associations. The mean latency of these differential responses was 154 +/- 44 (SD) ms. The firing of these neurons was shown to reflect a combination of the particular stimulus and the particular response associated by learning in the stimulus-response association task and could not be accounted for by the motor requirements of the task, nor wholly the stimulus aspects of the task, as demonstrated by testing their firing in related visual discrimination tasks. Responsive neurons were found throughout the hippocampal formation, but were particularly concentrated in the subicular complex and the CA3 subfield. These results show that single hippocampal neurons respond to combinations of the visual stimuli and the spatial responses with which they must become associated in conditional spatial response tasks and are consistent with the suggestion that part of the mechanism of this learning involves associations between visual stimuli and spatial responses learned by single hippocampal neurons.

Action Potentials↗

PubMed1989

Information processing in the taste system of primates.

1. Analysis of the activity of single neurones in the gustatory pathways in primates (cynomolgus monkeys) shows that the tuning of neurones to the four prototypical stimuli 1.0 mol l-1 glucose, 1.0 mol l-1 NaCl, 0.001 mol l-1 quinine-HCl and 0.01 mol l-1 HCl becomes sharper as information progresses through the taste system from the first central relay in the brainstem, the nucleus of the solitary tract, via the thalamus to the primary taste cortex in the frontal operculum and insula to reach a secondary cortical taste area in the caudolateral orbitofrontal cortex. 2. Feeding monkeys to satiety with glucose has no effect on the gustatory responses of neurones in the nucleus of the solitary tract, the frontal opercular taste cortex or the insular taste cortex, but decreases the magnitude of the neuronal responses of orbitofrontal cortex taste neurones which respond to glucose to zero. 3. The responses of orbitofrontal cortex taste neurones decrease to foods on which the monkey is fed to satiety, but continue to foods which have not just been eaten, that is, they reflect sensory-specific satiety, the phenomenon in which the pleasantness of the taste of a food and its acceptability, but not those of of other foods, is decreased by eating that food to satiety. 4. It has been found that the orbitofrontal cortex taste area or areas receives inputs from different modalities: single neurones with unimodal responses to taste (47%), olfactory (12%) and visual (10%) stimuli are found in close proximity to each other. Moreover, some single neurones show multimodal convergence, responding, for example, to taste and visual inputs (17%), taste and olfactory inputs (10%), and olfactory and visual inputs (4%). Some of these multimodal single neurones have corresponding sensitivities in the two modalities: they respond best to sweet tastes (e.g. 1 mol l-1 glucose), and respond more in a visual discrimination task to the visual stimulus which signifies sweet fruit juice than to that which signifies saline; or they respond to sweet taste and, in an olfactory discrimination task, to fruit juice odour. These results suggest that multimodal representations are formed in the orbitofrontal cortex secondary and related taste areas.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

PubMed1989

Hippocampal neurons in the monkey with activity related to the place in which a stimulus is shown.

In order to analyze the functions of the hippocampus in the primate, and to advance the understanding of amnesia, the activity of 994 single hippocampal neurons in the monkey was analyzed during the performance of a task known to be affected by hippocampal damage in which both an object, and its position in space, must be remembered. The serial multiple object-place memory task required a memory for the position on a video monitor in which a given object had appeared previously. It was found that 9.3% of neurons recorded in the hippocampus and hippocampal gyrus had spatial fields in this and related tasks, in that they responded whenever there was a stimulus in some but not in other positions on the screen. We found that 2.4% of the neurons responded to a combination of spatial information and information about the object seen, in that they responded more the first time a particular object was seen in any position. Six of these neurons were found that showed this combination even more clearly, in that, for example, they responded only to some positions and only if it was the first time that a particular stimulus had appeared there. It is concluded that there are neurons in the primate hippocampus which (1) respond to position in space and (2) in some cases combine information about stimuli and their position in space, responding to a stimulus only the first time it is seen in a position in space, for example. Thus, not only is spatial information processed by the primate hippocampus, but it can be combined with information about which stimuli have been seen before. The ability of the hippocampus to form such conjunctions may be an important property for its role in memory.

Analysis of Variance↗

PubMed1988

Neuronal activity of the supplementary motor area (SMA) during internally and externally triggered wrist movements.

The activity of neurones was recorded from the supplementary motor area (SMA) of monkeys while they were performing a discrete, arbitrary wrist movement. The cells responded similarly whether there was a triggering stimulus at the time of the movement or not. This experiment indicates that SMA neurones are active both in relation to externally triggered and internally initiated (voluntary) actions.

Animals↗

PubMed1988

The responsiveness of neurons in the insular gustatory cortex of the macaque monkey is independent of hunger.

(1) In order to determine whether the responsiveness of neurons in the insular gustatory cortex is influenced by hunger, neuronal activity was analysed in it while macaque monkeys (Macaca fascicularis) were fed to satiety. The responses of single neurons in the insular gustatory cortex to the protypical taste stimuli glucose, NaCl, HCl and quinine HCl, and to fruit juice, were measured before, while, and after the monkey was fed to satiety with glucose or fruit juice. (2) While behavior turned from avid acceptance to active rejection upon repletion, the responsiveness of the neurons to the stimulus array, including the satiating solution, was unmodified. (3) It is concluded that in the insular gustatory cortex, neuronal responses to gustatory stimuli are not influenced by the normal transition from hunger to satiety. This is in contrast to the responses of a population of neurons recorded in the hypothalamus, which only respond to the taste of food when the monkey is hungry. Thus the neurons in the insular gustatory cortex are involved in a motivation-independent analysis of gustatory stimuli, whereas the hypothalamic neurons may be more closely related to the influence of motivational state on behavioral responsiveness to gustatory stimuli.

Amygdala↗

PubMed1988

The responsiveness of neurones in the frontal opercular gustatory cortex of the macaque monkey is independent of hunger.

1. In order to determine whether the responsiveness of neurones in the primary gustatory cortex is influenced by hunger, the activity of neurones in the gustatory cortex in the frontal operculum was recorded while macaque monkeys (Macaca fascicularis) were fed to satiety. The responses of single neurones in the gustatory cortex to the prototypical taste stimuli glucose, NaCl, HCl and quinine hydrochloride, and to fruit juice, were measured before, while, and after the monkey was fed to satiety with glucose or fruit juice. 2. While behaviour turned from avid acceptance to active rejection upon repletion, the responsiveness of the neurones to the stimulus array, including the satiating solution, was unmodified. 3. It is concluded that in the gustatory cortex in the frontal operculum, neuronal responses to gustatory stimuli are not influenced by the normal transition from hunger to satiety. This is in contrast to the responses of a population of neurones recorded in the hypothalamus, which only occur to the taste of food when the monkey is hungry. Thus the neurones in the primary gustatory cortex are involved in a motivation-independent analysis of gustatory stimuli, whereas the hypothalamic neurones may be more closely related to the influence of motivational state on behavioural responsiveness to gustatory stimuli.

Action Potentials↗

PubMed1987

Responses of neurons in the inferior temporal cortex in short term and serial recognition memory tasks.

Gaffan and Weiskrantz (1980) and Mishkin (1982) have shown that lesions to the inferior temporal visual cortex can impair the performance of serial visual recognition memory tasks. In order to provide evidence on whether the inferior temporal visual cortex contains a mechanism which enables memory to span the intervening items in a serial recognition task, or whether the inferior temporal cortex is merely afferent to such recent memory mechanisms, we analysed the activity of single neurons in the inferior temporal visual cortex and the adjacent cortex in the superior temporal sulcus in both delayed match to sample and serial recognition memory tasks. In the serial recognition task, various numbers of stimuli intervened between the first and second presentations of a stimulus. A considerable proportion (64/264 or 26%) of visually responsive inferotemporal neurons showed a different response to the "novel" and "familiar" presentations of a stimulus in the serial recognition memory task, and often a corresponding difference in response between the sample and match presentations of a stimulus in the delayed match to sample task. For the majority of neurons this difference was not sustained across even one intervening stimulus in the serial recognition task, and no neurons bridged more than 2 intervening stimuli. These results show that neurons in the inferior temporal cortex have responses which would be useful for a short term visual memory for stimuli, but would not be useful in recency memory tasks in which more than one stimulus intervenes between the first and second presentations of a stimulus. In this investigation, neurons were recorded both in the cortex on the inferior temporal gyrus (commonly called inferior temporal visual cortex, and consisting of areas TE3, TE2 and TE1 of Seltzer and Pandya 1978), and in the cortex in the adjacent anterior part of the superior temporal sulcus, in which a number of different temporal cortical visual areas have now been described.

Animals↗

PubMed1987

The responses of neurons in the cortex in the superior temporal sulcus of the monkey to band-pass spatial frequency filtered faces.

There are neurons in the cortex in the anterior part of the superior temporal sulcus of the macaque monkey with visual responses which would be useful for face recognition (Rolls, 1984; Baylis et al., 1985). To analyze further the information which leads them to respond, their responses were measured to parametrically filtered stimuli. The responses of 48 such single neurons were measured to faces which were digitized and were bandpass spatial frequency filtered. The octave width bands were 2-4, 4-8, 8-16, 16-32, 32-64 and 64-128 cycles per image. It was found that the neurons could respond well to single octaves of the spatial frequencies normally present in faces, that the most effective bands were 4-8, 8-16 and 16-32 cycles per face (cpf), and that the bands 2-4 and 32-64 cpf were partly effective. In investigations of whether the responses of the neurons to an unfiltered face, and to low-pass and high-pass filtered images could be predicted by linear addition of their responses to each of the octave bands shown separately, it was found that the majority of the neurons were non-linear, and responded much less than predicted. It was also shown that this occurred even when the contrast was reduced to 0.25 of that normally present in a face, so that the result was not due just to a ceiling effect of the maximum firing rate. These results help to define parametrically the aspects of the information normally present in a face which are sufficient to produce responses of these neurons to them, and show that linear operations cannot account for information processing in this part of the visual system.

Action Potentials↗

PubMed1987

Functional subdivisions of the temporal lobe neocortex.

In order to gather evidence on functional subdivisions of the temporal lobe neocortex of the primate, the activity of more than 2600 single neurons was recorded in 10 myelo- and cytoarchitecturally defined subdivisions of the cortex in the superior temporal sulcus (STS) and inferior temporal gyrus of the anterior part of the temporal lobe of 5 hemispheres of 3 macaque monkeys. First, convergence of different modalities into each area was investigated. Areas TS and TAa, in the upper part of this region, were found to receive visual as well as auditory inputs. Areas TPO, PGa, and IPa, in the depths of the STS, received visual, auditory, and somatosensory inputs. Areas TEa, TEm, TE3, TE2, and TE1, which extend from the ventral bank of the STS through the inferior temporal gyrus, were primarily unimodal visual areas. Second, of the cells with visual responses, it was found that some neurons in areas TS-IPa could be activated only by moving visual stimuli, whereas the great majority of neurons in areas TEa-TE1 could be activated by stationary visual stimuli. Third, it was found that there were few sharply discriminating visual neurons in areas TS and TAa; of the sharply discriminating visual neurons in other areas, however, neurons that responded primarily to faces were found predominantly in areas TPO, TEa, and TEm (in which they represented 20% of the neurons with visual responses); neurons that were tuned to relatively simple visual stimuli such as sine-wave gratings, color, or simple shapes were relatively common in areas TEa, TEm, and TE3; and neurons that responded only to complex visual stimuli were common in areas IPa, TEa, TEm, and TE3. These findings show inter alia that areas TPO, PGa, and IPa are multimodal, that the inferior temporal gyrus areas are primarily unimodal, that there are areas in the cortex in the anterior and dorsal part of the STS that are specialized for the analysis of moving visual stimuli, that neurons responsive primarily to faces are found predominantly in areas TPO, TEa, and TEm, and that architectural subdivisions of the temporal lobe cortex are related to neuronal response properties.

Action Potentials↗

PubMed1986

Sensory-specific satiety: food-specific reduction in responsiveness of ventral forebrain neurons after feeding in the monkey.

It has been shown previously that some neurons in the lateral hypothalamus and substantia innominata respond to the sight of food, others to the taste of food, and others to the sight or taste of food, in the hungry monkey. It is shown here that feeding to satiety decreases the responses of hypothalamic neurons to the sight and/or taste of a food on which the monkey has been satiated, but leaves the responses of the same neurons to other foods on which the monkey has not been satiated relatively unchanged. This suggests that the responses of these neurons in the ventral forebrain are related to sensory-specific satiety, an important phenomenon which regulates food intake. In sensory-specific satiety, the pleasantness of the sight or taste of a food becomes less after it is eaten to satiety, whereas the pleasantness of the sight or taste of other foods which have not been eaten is much less changed; correspondingly, food intake is greater if foods which have not already been eaten to satiety are offered.

Animals↗

PubMed1986

Size and contrast have only small effects on the responses to faces of neurons in the cortex of the superior temporal sulcus of the monkey.

There is a population of neurons in the cortex in the middle and anterior part of the superior temporal sulcus (STS) of the monkey with responses which are selective for faces. To investigate whether the responses of these neurons show some of the perceptual properties of face recognition such as tolerance to changes in the size and contrast of the face, the effects of alteration of the size and contrast of an effective face stimulus on the responses of these neurons were analysed quantitatively in macaque monkeys. First, it was shown that the majority of these neurons had responses which were relatively invariant with respect to the size of the stimulus. The median size change tolerated with a response of greater than half the maximal response was 12 times. Second, it was found that for a few of these neurons, the size of the face did affect the neuronal response. For most of these neurons, it was found that when the size of the image and its distance were altered, the neuronal response was related to the retinal angle subtended by the image. But for four neurons the absolute size of the image determined the magnitude of the neuronal response, independently of the distance of the image. Thus these four neurons showed size constancy. It is suggested that these neurons would be useful as part of a face recognition system, because only objects in a certain absolute size range should normally be classified as faces. Third, the responses of the neurons were relatively invariant with respect to the contrast of the face. The mean contrast at which the neurons still responded with more than half the maximal response was 0.26. Fourth, the responses of the neurons were relatively invariant with respect to the sign of the contrast of the face, that is the neurons responded to negative as well as to positive images of faces. Fifth, the neurons typically responded to a face when the information in it had been reduced from 3D to a 2D representation in gray on a monitor, with a response which was on average 0.5 that to a real face. These results show that the responses of these neurons have some of the invariant properties with respect to size and contrast alteration shown by face perception, and show that their processing is at a level which would be useful in face recognition.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

PubMed1986

Gustatory responses in the nucleus tractus solitarius of the alert cynomolgus monkey.

Multiunit and single neuron responses to taste stimuli in the nucleus tractus solitarius (NTS) of alert cynomolgus monkeys were analyzed. Intensity-response functions, including neural thresholds to glucose and quinine HCl, agreed well with psychophysical reports, implying that the cynomolgus monkey and human share the same dynamic range of sensitivity to prototypical taste stimuli. The NTS is chemotopically organized: neurons most responsive to HCl are more common in the posterior gustatory area, whereas those most responsive to glucose and NaCl are located in the anterior NTS. Responsiveness to quinine is more widely distributed but tends toward the anterior. Efforts were made to determine if neurons could be divided into a discrete number of types, as determined by their sensitivities to the prototypical stimuli. The clearest distinction was between those that did or did not respond well to HCl. Beyond this, neuronal categories were not obvious. Individual neurons were quite broadly sensitive to our stimulus array, so that, for the typical NTS cell, no one of the four prototypes evoked a majority of the discharges. This extreme breadth of tuning suggests that taste-quality information in the monkey might be incorporated in relative discharge rates across the neuron population. Correlations among patterns of activity to the four prototypical stimuli indicated that only HCl and quinine HCl have closely related taste qualities.

Afferent Pathways↗

PubMed1986

Gustatory responses in the frontal opercular cortex of the alert cynomolgus monkey.

The responses of 165 single taste neurons in the anterior operculum of the alert cynomolgus monkey were analyzed. Chemicals were deionized water, blackcurrant juice, and the four basic taste stimuli: glucose, NaCl, HCl, and quinine HCl. Taste-evoked responses could be recorded from an opercular region that measured approximately 4.0 mm in its anteroposterior extent, 2.0 mm mediolaterally, and 3.0 mm dorsoventrally. Within this area, taste-responsive neurons were sparsely distributed such that multiunit activity was rarely encountered and neuronal isolation was readily achieved. Intensity-response functions were determined for nine cells. In each case, the lowest concentration of the dynamic response range conformed well to human electrophysiological and psychophysical thresholds for the basic taste stimuli. There was some evidence of chemotopic organization. Cells that responded best to glucose tended to be distributed toward the anterior operculum, whereas most acid-sensitive neurons were located more posteriorly. The proportion of cells responding best to NaCl peaked in the middle of the area, whereas quinine sensitivity was rather evenly distributed throughout. Opercular neurons in the monkey showed moderate breadth of sensitivity compared with taste cells of other species and at other synaptic levels. A breadth-of-tuning coefficient was calculated for each neuron. This is a metric that can range from 0.0 for a cell that responds specifically to only one of the four basic stimuli to 1.0 for one that responds equally to all four stimuli. The mean coefficient for 165 cells in the operculum was 0.67 (range = 0.12-0.99). Efforts were made to determine whether neurons could be divided into a discrete number of types, as defined by their responsiveness to the stimulus array used here. It was concluded that most taste cells may be assigned to a small number of groups, each of which is statistically independent of the others, but within which the constituent neurons are not identical. An analysis of taste quality indicated that the sweet and salty stimuli evoked patterns of activity that were significantly intercorrelated. Similarly, patterns representing HCl, quinine HCl, and water were related.

Animals↗

PubMed1985

Satiety does not affect gustatory activity in the nucleus of the solitary tract of the alert monkey.

Feeding to satiety decreases the acceptability of the taste of food. In order to determine whether the responsiveness of gustatory neurons in the nucleus tractus solitarius (NTS) is influenced by hunger, neural activity in the NTS was analyzed while monkeys were fed to satiety. Gustatory neural activity to glucose, fruit juice, NaCl, HCl and quinine HCl was measured before, while and after the monkey was fed to satiety with glucose, fruit juice or sucrose. While behavior turned from avid acceptance to active rejection upon repletion, the responsiveness of NTS neurons to the stimulus array, including the satiating solution, was unmodified. It is concluded that at the first central synapse of the taste system of the primate, neural responsiveness is not influenced by the normal transition from hunger to satiety. This is in contrast to the responses of a population of neurons recorded in the hypothalamus, which only occur to the taste of food when the monkey is hungry. Thus, NTS gustatory activity appears to occur independently of normal hunger and satiety, whereas hypothalamic neuronal activity is more closely related to the influence of motivational state on behavioral responsiveness to gustatory stimuli.

Animals↗

PubMed1985

Selectivity between faces in the responses of a population of neurons in the cortex in the superior temporal sulcus of the monkey.

There is a population of neurons in the cortex in the middle and anterior part of the superior temporal sulcus (STS) of the monkey with responses which are selective for faces. If, consistent with the effects of damage to the temporal lobe, these neurons are involved in face recognition or in making appropriate social responses to different individuals, then it might be expected that at least some of these neurons might respond differently to different faces. To investigate whether at least some of these neurons do respond differently to different faces, their responses were measured to a standard set of faces, presented in random sequence using a video framestore. It was found that a considerable proportion of the neurons with face-selective responses tested (34/44 or 77%) responded differently to different faces, as shown by analyses of variance. An index of the discriminability of the most and least effective face stimulus (d') ranged between 0.2 and 5.0 for the different neurons. Although these neurons often responded differently to different faces, they did not usually respond to only one of the faces in the set, so that information that a particular face had been shown was present across an ensemble of neurons, rather than in the responses of an individual neuron. These findings indicate that the responses of these neurons would be useful in providing information on which different behavioral responses made to different faces could be based. These neurons could thus be filters, the output of which could be used for recognition of different individuals and in emotional responses made to different individuals.

Animals↗

PubMed1985

Long-term sensory-specific satiety: evidence from an Ethiopian refugee camp.

The reduction in appetite which occurs during a meal is partly specific to the foods which have been eaten earlier in a meal. This has been called "sensory-specific satiety." In the experiment described here, a long-term form of sensory-specific satiety has been demonstrated. Refugees in an Ethiopian refugee camp reported that the taste of 3 foods which they had been eating for approximately 6 months was less pleasant than that of 3 new foods, whereas refugees who had been eating the regular diet for only two days found its taste as pleasant as that of the different foods. This long-term sensory-specific satiety may have nutritional implications when only a limited variety of food is available, as may occur in refugee camps. It will be useful to determine the extent to which this long-term sensory-specific satiety can be prevented by provision of perhaps even a limited range of spices, flavorings or foods.

Adolescent↗

PubMed1985

Role of low and high spatial frequencies in the face-selective responses of neurons in the cortex in the superior temporal sulcus in the monkey.

There are neurons in the cortex in the anterior part of the superior temporal sulcus of the macaque monkey with visual responses selective for faces. One aim of the present study was to analyze further the information which leads them to respond, by measuring their responses to parametrically filtered stimuli. The responses of 32 such single neurons were measured to faces which were digitized, lowpass filtered at spatial frequencies of 2, 4, 8,...128 cycles/face, highpass filtered at frequencies of 4, 8,...64 cycles/face, and presented in random sequence using a video framestore. It was found that many of the neurons could respond to blurred images of faces, with a mean frequency at half-maximum amplitude of the neuronal response to the series of lowpass filtered images of faces of 3.3 cycles/face. Almost all the neurons had lowpass cutoff frequencies defined in this way below 8 cycles/face. Many of the neurons could also respond to images of faces in which the only information remaining was a limited amount of high spatial frequency edge information. The mean frequency at half-maximum amplitude of the neuronal response to the series of highpass filtered images of faces was 29.7 cycles/face. Almost all the neurons had highpass cutoff frequencies above 8 cycles/face. Thus, many of the neurons could respond to a lowpass and a highpass filtered image of a face even when these had no spatial frequencies in common. The mean separation between the lowpass and highpass cutoff frequencies was 3.2 octaves. For comparison, face recognition in man can be performed with images which contain only information up to 8 cycles/face, or with highpass filtered images which contain only information down to 8 cycles/face. The response of the neurons was not always a smooth function of frequency, but could decrease as higher frequencies were included in the lowpass filtered images of faces, or as low frequencies were included in the highpass filtered images of faces. This indicates that information in certain frequency bands was able to inhibit these neurons. This was particularly likely to occur for the non-optimal face stimulus for a given neuron, indicating that the selectivity of these neurons to different faces was a combination of the excitation produced by some information in faces and inhibition produced by other.

Action Potentials↗