CiteULike: klouie's neurophysiology

A Neural Correlate of Motivational Conflict in the Superior Colliculus of the Macaque

Masaki Isoda — 2008-07-03T16:23:32-00:00

J Neurophysiol (2 July 2008), 90275.2008.

Behavior is controlled by both external instructions and internal motives, but the actions demanded by each may be different. A common consequence of such a conflict is a delay in decision making and subsequent motor responses. It is unknown, however, what neural mechanisms underlie motivational conflict and associated response delay. To answer this question, we recorded single-neuron activity in the superior colliculus (SC) as macaque monkeys performed a visually guided, asymmetrically rewarded saccade task. A peripheral spot of light at one of two opposing positions was illuminated to indicate a saccade target. In a given block of trials, one position was associated with a big reward and the other with a small reward. The big-reward position was alternated across blocks. Behavioral analyses revealed that small-reward trials created a conflict between the instructed saccade to one position and the internally motivated, yet invalid saccade to the opposite position. We found that movement neurons in the SC temporally exhibited bursting activity after the appearance of the small-reward target opposite their movement field. This transient activity predicted the amount of response delay for upcoming saccades. Our data suggest that motivational conflict activates movement neurons in both colliculi, thereby delaying saccade initiation through intercollicular inhibitory interactions. 10.1152/jn.90275.2008

Simultaneous encoding of multiple potential reach directions in dorsal premotor cortex.

P Cisek — 2005-11-25T05:22:20-00:00

J Neurophysiol, Vol. 87, No. 2. (February 2002), pp. 1149-1154.

We present evidence that the primate brain can simultaneously generate discrete directional signals related to multiple alternative reaching actions before making a decision between them. A monkey performed a task in which the correct target for a reaching movement was specified during two consecutive instructed-delay periods. First, two potential targets were presented; and second, a nonspatial cue identified one of them as the correct movement target. During the first period, two directional signals coexisted in the activity of cells in dorsal premotor cortex (PMd), oriented toward the two potential targets. During the second period, one of these disappeared and the remaining signal predicted the monkey's response choice. These results suggest that, when faced with multiple salient opportunities for reaching, the primate brain performs sensorimotor transformations in parallel to begin planning several reaching movements simultaneously before selecting one for overt execution.

Neural correlates of reaching decisions in dorsal premotor cortex: specification of multiple direction choices and final selection of action.

P Cisek — 2007-04-08T21:09:33-00:00

Neuron, Vol. 45, No. 5. (3 March 2005), pp. 801-814.

We show that while a primate chooses between two reaching actions, its motor system first represents both options and later reflects selection between them. When two potential targets appeared, many (43%) task-related, directionally tuned cells in dorsal premotor cortex (PMd) discharged if one of the targets was near their preferred direction. At the population level, this generated two simultaneous sustained directional signals corresponding to the current reach options. After a subsequent nonspatial cue identified the correct target, the corresponding directional signal increased, and the signal for the rejected target was suppressed. The PMd population reliably predicted the monkey's response choice, including errors. This supports a planning model in which multiple reach options are initially specified and then gradually eliminated in a competition for overt execution, as more information accumulates.

Decision-making with multiple alternatives

Anne Churchland — 2008-05-27T17:05:46-00:00

Nat Neurosci, Vol. 11, No. 6. (June 2008), pp. 693-702.

Electrical microstimulation thresholds for behavioral detection and saccades in monkey frontal eye fields

Dona Murphey — 2008-05-27T15:49:43-00:00

Proceedings of the National Academy of Sciences (13 May 2008), 0710820105.

The frontal eye field (FEF) is involved in the transformation of visual signals into saccadic eye movements. Although it is often considered an oculomotor structure, several lines of evidence suggest that the FEF also contributes to visual perception and attention. To better understand the range of behaviors to which the FEF can contribute, we tested whether monkeys could detect activation of their FEF by electrical microstimulation with currents below those that cause eye movements. We found that stimulation of FEF neurons could almost always be detected at levels below those needed to generate saccades and that the electrical current needed for detection was highly correlated with that needed to generate a saccade. This relationship between detection and saccade thresholds can be explained if FEF neurons represent preparation to make particular saccades and subjects can be aware of such preparations without acting on them when the representation is not strong. 10.1073/pnas.0710820105

Reward-Dependent Modulation of Neuronal Activity in the Primate Dorsal Raphe Nucleus

Kae Nakamura — 2008-05-14T17:09:11-00:00

J. Neurosci., Vol. 28, No. 20. (14 May 2008), pp. 5331-5343.

The dopamine system has been thought to play a central role in guiding behavior based on rewards. Recent pharmacological studies suggest that another monoamine neurotransmitter, serotonin, is also involved in reward processing. To elucidate the functional relationship between serotonin neurons and dopamine neurons, we performed single-unit recording in the dorsal raphe nucleus (DRN), a major source of serotonin, and the substantia nigra pars compacta, a major source of dopamine, while monkeys performed saccade tasks in which the position of the target indicated the size of an upcoming reward. After target onset, but before reward delivery, the activity of many DRN neurons was modulated tonically by the expected reward size with either large- or small-reward preference, whereas putative dopamine neurons had phasic responses and only preferred large rewards. After reward delivery, the activity of DRN neurons was modulated tonically by the received reward size with either large- or small-reward preference, whereas the activity of dopamine neurons was not modulated except after the unexpected reversal of the position-reward contingency. Thus, DRN neurons encode the expected and received rewards, whereas dopamine neurons encode the difference between the expected and received rewards. These results suggest that the DRN, probably including serotonin neurons, signals the reward value associated with the current behavior. 10.1523/JNEUROSCI.0021-08.2008

Neural Activity in the Frontal Eye Fields Modulated by the Number of Alternatives in Target Choice

Kyoung-Min Lee — 2008-05-22T10:00:57-00:00

J. Neurosci., Vol. 28, No. 9. (27 February 2008), pp. 2242-2251.

Selection of identical responses may not use the same neural mechanisms when the number of alternatives (NA) for the selection changes, as suggested by Hick's law. For elucidating the choice mechanisms, frontal eye field (FEF) neurons were monitored during a color-to-location choice saccade task as the number of potential targets was varied. Visual responses to alternative targets decreased as NA increased, whereas perisaccade activities increased with NA. These modulations of FEF activities seem closely related to the choice process because the activity enhancements coincided with the timing of target selection, and the neural modulation was greater as NA increased, features expected of neural correlates for a choice process from the perspective of Hick's law. Our current observations suggest two novel notions of FEF neuronal behavior that have not been reported previously: (1) cells called "phasic visual" that do not discharge in the perisaccade interval in a delayed-saccade paradigm show such activity in a choice response task at the time of the saccade; and (2) the activity in FEF visuomotor cells display an inverse relationship between perisaccadic activity and the time of saccade triggering with higher levels of activity leading to longer saccade reaction times. These findings support the area's involvement in sensory-motor translation for target selection through coactivation and competitive interaction of neural populations that code for alternative action sets. 10.1523/JNEUROSCI.3596-07.2008

Expectation of reward modulates cognitive signals in the basal ganglia.

R Kawagoe — 2007-01-29T21:56:50-00:00

Nat Neurosci, Vol. 1, No. 5. (September 1998), pp. 411-416.

Action is controlled by both motivation and cognition. The basal ganglia may be the site where these kinds of information meet. Using a memory-guided saccade task with an asymmetric reward schedule, we show that visual and memory responses of caudate neurons are modulated by expectation of reward so profoundly that a neuron's preferred direction often changed with the change in the rewarded direction. The subsequent saccade to the target was earlier and faster for the rewarded direction. Our results indicate that the caudate contributes to the determination of oculomotor outputs by connecting motivational values (for example, expectation of reward) to visual information.

Reward-dependent gain and bias of visual responses in primate superior colliculus.

T Ikeda — 2008-05-06T22:49:25-00:00

Neuron, Vol. 39, No. 4. (14 August 2003), pp. 693-700.

Eye movements are often influenced by expectation of reward. Using a memory-guided saccade task with an asymmetric reward schedule, we show that visual responses of monkey SC neurons increase when the visual stimulus indicates an upcoming reward. The increase occurred in two distinct manners: (1) reactively, as an increase in the gain of the visual response when the stimulus indicated an upcoming reward; (2) proactively, as an increase in anticipatory activity when reward was expected in the neuron's response field. These effects were observed mostly in saccade-related SC neurons in the deeper layer which would receive inputs from the cortical eye fields and the basal ganglia. These results, together with recent findings, suggest that the gain modulation may be determined by the inputs from both the cortical eye fields and the basal ganglia, whereas the anticipatory bias may be derived mainly from the basal ganglia.

Integration of exogenous input into a dynamic salience map revealed by perturbing attention.

PF Balan — 2006-09-14T16:39:44-00:00

J Neurosci, Vol. 26, No. 36. (6 September 2006), pp. 9239-9249.

Although it is widely accepted that exogenous and voluntary factors jointly determine the locus of attention, the rules governing the integration of these factors are poorly understood. We investigated neural responses in the lateral intraparietal area (LIP) to transient, distracting visual perturbations presented during task performance. Monkeys performed a covert search task in which they discriminated the orientation of a target embedded among distractors, and brief visual perturbations were presented at various moments and locations during task performance. LIP neurons responded to perturbations consisting of the appearance of new objects, as well as to abrupt changes in the color, luminance, or position of existing objects. The LIP response correlated with the bottom-up behavioral effects of different perturbation types. In addition, neurons showed two types of top-down modulations. One modulation was a context-specific multiplicative gain that affected perturbation, target, and distractor activity in a spatially nonspecific manner. Gain was higher in blocks of trials in which perturbations directly marked target location than in blocks in which they invariably appeared opposite the target, thus encoding a behavioral context defined by the statistical contingency between target and perturbation location. A second modulation reflected local competitive interactions with search-related activity, resulting in the converse effect: weaker perturbation-evoked responses if perturbations appeared at the location of the target than if they appeared opposite the target. Thus, LIP encodes an abstract dimension of salience, which is shaped by local and global top-down mechanisms. These interacting mechanisms regulate responsiveness to external input as a function of behavioral context and momentary task demands.

Perceptual and motor processing stages identified in the activity of macaque frontal eye field neurons during visual search.

KG Thompson — 2008-05-05T20:22:50-00:00

Journal of neurophysiology, Vol. 76, No. 6. (December 1996), pp. 4040-4055.

1. The latency between the appearance of a popout search display and the eye movement to the oddball target of the display varies from trial to trial in both humans and monkeys. The source of the delay and variability of reaction time is unknown but has been attributed to as yet poorly defined decision processes. 2. We recorded neural activity in the frontal eye field (FEF), an area regarded as playing a central role in producing purposeful eye movements, of monkeys (Macaca mulatta) performing a popout visual search task. Eighty-four neurons with visually evoked activity were analyzed. Twelve of these neurons had a phasic response associated with the presentation of the visual stimulus. The remaining neurons had more tonic responses that persisted through the saccade. Many of the neurons with more tonic responses resembled visuomovement cells in that they had activity that increased before a saccade into their response field. 3. The visual response latencies of FEF neurons were determined with the use of a Poisson spike train analysis. The mean visual latency was 67 ms (minimum = 35 ms, maximum = 138 ms). The visual response latencies to the target presented alone, to the target presented with distractors, or to the distractors did not differ significantly. 4. The initial visual activation of FEF neurons does not discriminate the target from the distractors of a popout visual search stimulus array, but the activity evolves to a state that discriminates whether the target of the search display is within the receptive field. We tested the hypothesis that the source of variability of saccade latency is the time taken by neurons involved in saccade programming to select the target for the gaze shift. 5. With the use of an analysis adapted from signal detection theory, we determined when the activity of single FEF neurons can reliably indicate whether the target or distractors are present within their response fields. The time of target discrimination partitions the reaction time into a perceptual stage in which target discrimination takes place, and a motor stage in which saccade programming and generation take place. The time of target discrimination occurred most often between 120 and 150 ms after stimulus presentation. 6. We analyzed the time course of target discrimination in the activity of single cells after separating trials into short, medium, and long saccade latency groups. Saccade latency was not correlated with the duration of the perceptual stage but was correlated with the duration of the motor stage. This result is inconsistent with the hypothesis that the time taken for target discrimination, as indexed by FEF neurons, accounts for the wide variability in the time of movement initiation. 7. We conclude that the variability observed in saccade latencies during a simple visual search task is largely due to postperceptual motor processing following target discrimination. Signatures of both perceptual and postperceptual processing are evident in FEF. Procrastination in the output stage may prevent stereotypical behavior that would be maladaptive in a changing environment.

Saccade and blinking evoked by microstimulation of the posterior parietal association cortex of the monkey.

H Shibutani — 2008-05-05T18:35:10-00:00

Experimental brain research. Experimentelle Hirnforschung. Expérimentation cérébrale, Vol. 55, No. 1. (1984), pp. 1-8.

Electrical stimulation with microelectrodes of the posterior parietal association cortex in alert behaving monkeys elicited saccadic eye movements and blinking. The sites in which saccades were elicited by electrical stimulation were concentrated in the anteromedial part of area 7a, especially in the posterior bank of the intraparietal sulcus, in a region which sends efferent projections to the frontal eye field and the superior colliculus, but they were also found in the posterolateral part of area 7a. Compared with the frontal eye fields and the superior colliculus, the threshold current for eliciting saccades was relatively high, on the average 86 microA. Moreover, the elicitation of saccade was inconsistent even with suprathreshold stimulation and suppressed during visual fixation. Latencies of the saccades were relatively long, on the average 50ms; they were longer in the posterolateral part than in the anteromedial part. Direction and amplitude of evoked saccades depended on the site of stimulation, but was independent of eye position in most cases. However, "goal-directed" saccades which depended on initial eye position were elicited in three penetrations in the posterolateral part of area 7a. The threshold of mainly in the lateral part of area 7a. The threshold of blinking was 70 microA and the latency was 50 ms on the average. In contrast to saccades, blinking was elicited constantly with each stimulus even during attentive fixation. We occasionally recorded single unit activity at the site of stimulation with the same electrodes. More than half of the units recorded at the site of blinking responded to approaching visual stimulus.(ABSTRACT TRUNCATED AT 250 WORDS)

A neural basis for visual search in inferior temporal cortex.

L Chelazzi — 2007-05-17T14:56:32-00:00

Nature, Vol. 363, No. 6427. (27 May 1993), pp. 345-347.

We often search for a face in a crowd or for a particular object in a cluttered environment. In this type of visual search, memory interacts with attention: the mediating neural mechanisms should include a stored representation of the object and a means for selecting that object from among others in the scene. Here we test whether neurons in inferior temporal cortex, an area known to be important for high-level visual processing, might provide these components. Monkeys were presented with a complex picture (the cue) to hold in memory during a delay period. The cue initiated activity that persisted through the delay among the neurons that were tuned to its features. The monkeys were then given 2-5 choice pictures and were required to make an eye movement to the one (the target) that matched the cue. About 90-120 milliseconds before the onset of the eye movement to the target, responses to non-targets were suppressed and the neuronal response was dominated by the target. The results suggest that inferior temporal cortex is involved in selecting the objects to which we attend and foveate.

Covert orienting of attention in macaques. II. Contributions of parietal cortex.

DL Robinson — 2008-04-30T18:11:53-00:00

Journal of neurophysiology, Vol. 74, No. 2. (August 1995), pp. 698-712.

1. To understand some of the contributions of parietal cortex to the dynamics of visual spatial attention, we recorded from cortical cells of monkeys performing attentional tasks. We studied 484 neurons in the intraparietal sulcus and adjacent gyral tissue of two monkeys. We measured phasic responses to peripheral visual stimuli while the monkeys attended toward or away from the stimuli or when attention was not controlled. Neurons were tested while the monkeys gazed at a spot of light (simple fixation task), actively attended to a foveal target (foveal attention task), performed a reaction time task (cued reaction time task), made saccadic eye movements to visual targets (saccade task), or responded to a repetitious peripheral target (probability task). 2. In a previous paper we demonstrated that monkeys, like humans, responded more quickly to visual targets when the targets followed briefly flashed visual cues (validly cued targets) (Bowman et al. 1993). It has been hypothesized that the cue attracts attention to its locus and results in faster reaction times (Posner 1980). In the present physiological studies, visual cues consistently excited these neurons when they were flashed in the receptive field. Such activity might signal a shift of attention. Visual targets that fell within the receptive field and that immediately followed the cue evoked relatively weak responses. This response was due to a relative refractory period. 3. Next we tested attentional processes in these tasks that were independent of the visual response to the cue. We placed the cue outside of the receptive field and the target within the receptive field. We found that 23% of these cells had a significant decrease in their firing rate to validly cued targets in their receptive fields under these conditions. Strong responses were evoked by the same target when the cue was flashed in the opposite hemifield (invalidly cued targets). Thus this group of neurons responded best when attention was directed toward the opposite hemifield. 4. For another group of parietal cells (13%) there was an enhanced response to targets in the visual receptive field when the cue was in the same hemifield. For the remaining 64% of the cells there was no significant modulation in this task. 5. The cued reaction time task involved exogenous control of attention; the sensory cue gave spatial and temporal direction to attention. We used several other tasks to test for endogenous control of attention.(ABSTRACT TRUNCATED AT 400 WORDS)

Response of neurons in the lateral intraparietal area to a distractor flashed during the delay period of a memory-guided saccade.

KD Powell — 2008-04-30T14:51:07-00:00

Journal of neurophysiology, Vol. 84, No. 1. (July 2000), pp. 301-310.

Recent experiments raised the possibility that the lateral intraparietal area (LIP) might be specialized for saccade planning. If this was true, one would expect a decreased sensitivity to irrelevant visual stimuli appearing late in the delay period of a memory-guided delayed-saccade task to a target outside the neurons' receptive fields. We trained two monkeys to perform a standard memory-guided delayed-saccade task and a distractor task in which a stimulus flashed for 200 ms at a predictable time 300-100 ms before the end of the delay period. We used two locations, one in the most active part of the receptive field and another well outside the receptive field. We used six kinds of trials randomly intermixed: simple delayed-saccade trials into or away from the receptive field and distractor trials with saccade target and distractor both in the receptive field, both out of the receptive field, or one at each location. This enabled us to study the response to the distractor as a function of the monkey's preparation of a memory-guided delayed-saccade task. We had assumed that the preparation of a saccade away from the receptive field would result in an attenuation of the response to the distractor, i.e., a distractor at the location of the saccade goal would evoke a greater response than when it appeared at a location far from the saccade goal. Instead we found that neurons exhibited either a normal or an enhanced visual response to the distractor during the memory period when the target flashed outside the receptive field. When the distractor flashed at the location of the saccade target, the response to the distractor was either unchanged or diminished. The response to a distractor away from the target location of a memory-guided saccade was even greater than the response to the same target when it was the target for the memory-guided saccade task. Immediate presaccadic activity did not distinguish between a saccade to the receptive field when there was no distractor and a distractor in the receptive field when the monkey made a saccade elsewhere. Nonetheless the distractor had no significant effect on the saccade latency, accuracy, or velocity despite the brisk response it evoked immediately before the saccade. We suggest that these results are inconsistent with a role for LIP in the specific generation of saccades, but they are consistent with a role for LIP in the generation of visual attention.

Neural Correlates of Attention and Distractibility in the Lateral Intraparietal Area

James Bisley — 2006-09-21T14:48:31-00:00

J Neurophysiol, Vol. 95, No. 3. (1 March 2006), pp. 1696-1717.

We examined the activity of neurons in the lateral intraparietal area (LIP) during a task in which we measured attention in the monkey, using an advantage in contrast sensitivity as our definition of attention. The animals planned a memory-guided saccade but made or canceled it depending on the orientation of a briefly flashed probe stimulus. We measured the monkeys' contrast sensitivity by varying the contrast of the probe. Both subjects had better thresholds at the goal of the saccade than elsewhere. If a task-irrelevant distractor flashed elsewhere in the visual field, the attentional advantage transiently shifted to that site. The population response in LIP correlated with the allocation of attention; the attentional advantage lay at the location in the visual field whose representation in LIP had the greatest activity when the probe appeared. During a brief period in which there were two equally active regions in LIP, there was no attentional advantage at either location. This time, the crossing point, differed in the two animals, proving a strong correlation between the activity and behavior. The crossing point of each neuron depended on the relationship of three parameters: the visual response to the distractor, the saccade-related delay activity, and the rate of decay of the transient response to the distractor. Thus the time at which attention lingers on a distractor is set by the mechanism underlying these three biophysical properties. Finally, we showed that for a brief time LIP neurons showed a stronger response to signal canceling the planned saccade than to the confirmation signal. 10.1152/jn.00848.2005

One-Dimensional Dynamics of Attention and Decision Making in LIP

Surya Ganguli — 2008-04-14T13:04:51-00:00

Neuron, Vol. 58, No. 1. (10 April 2008), pp. 15-25.

Summary Where we allocate our visual spatial attention depends upon a continual competition between internally generated goals and external distractions. Recently it was shown that single neurons in the macaque lateral intraparietal area (LIP) can predict the amount of time a distractor can shift the locus of spatial attention away from a goal. We propose that this remarkable dynamical correspondence between single neurons and attention can be explained by a network model in which generically high-dimensional firing-rate vectors rapidly decay to a single mode. We find direct experimental evidence for this model, not only in the original attentional task, but also in a very different task involving perceptual decision making. These results confirm a theoretical prediction that slowly varying activity patterns are proportional to spontaneous activity, pose constraints on models of persistent activity, and suggest a network mechanism for the emergence of robust behavioral timing from heterogeneous neuronal populations.

Activity changes in early visual cortex reflect monkeys' percepts during binocular rivalry.

DA Leopold — 2008-04-01T18:52:38-00:00

Nature, Vol. 379, No. 6565. (8 February 1996), pp. 549-553.

When the two eyes view dissimilar images, we experience binocular rivalry, in which one eye's view dominates for several seconds and is then replaced by that of the other eye. What causes these perceptual changes in the absence of any change in the stimulus? We showed previously that some neurons in monkey cortical area MT show changes in activity during motion rivalry that reflect the perceived direction of motion. To determine whether perception-related modulation of activity occurs in other visual cortical areas, we recorded from individual neurons in V1, V2 and V4 while monkeys reported the perceived orientation of rival gratings of two orthogonal orientations. Many cells, particularly in V4, showed patterns of activity that correlated with the perceptual dominance and suppression of one stimulus. The majority were orientation-selective and could be driven equally well from either eye. It has been previously suggested that binocular rivalry involves reciprocal inhibition between monocular neurons within V1 (for example, see ref. 4), but our results do not support this view; rather, we propose that binocular rivalry arises through interactions between binocular neurons at several levels in the visual pathways, and that similar mechanisms may underlie other multistable perceptual states that occur when viewing ambiguous images.

Perceptually Bistable Three-Dimensional Figures Evoke High Choice Probabilities in Cortical Area MT

Jonathan Dodd — 2008-04-01T18:35:27-00:00

J. Neurosci., Vol. 21, No. 13. (1 July 2001), pp. 4809-4821.

The role of the primate middle temporal area (MT) in depth perception was examined by considering the trial-to-trial correlations between neuronal activity and reported depth sensations. A set of moving random dots portrayed a cylinder rotating about its principal axis. In this structure-from-motion stimulus, the direction of rotation is ambiguous and the resulting percept undergoes spontaneous fluctuations. The stimulus can be rendered unambiguous by the addition of binocular disparities. We trained monkeys to report the direction of rotation in a set of these stimuli, one of which had zero disparity. Many disparity-selective neurons in area MT are selective for the direction of rotation defined by disparity. Across repeated presentations of the ambiguous (zero-disparity) stimulus, there was a correlation between neuronal firing and the reported direction of rotation, as found by Bradley et al. (1998). Quantification of this effect using choice probabilities (Britten et al., 1996) allowed us to demonstrate that the correlation cannot be explained by eye movements, behavioral biases, or attention to spatial location. MT neurons therefore appear to be involved in the perceptual decision process. The mean choice probability (0.67) was substantially larger than that reported for MT neurons in a direction discrimination task (Britten et al., 1996). This implies that MT neurons make a different contribution to the two tasks. For the depth task, either the pool of neurons used is smaller or the correlation between neurons in the pool is larger.

Bounded Integration in Parietal Cortex Underlies Decisions Even When Viewing Duration Is Dictated by the Environment

Roozbeh Kiani — 2008-03-19T18:16:47-00:00

J. Neurosci., Vol. 28, No. 12. (19 March 2008), pp. 3017-3029.

Decisions about sensory stimuli are often based on an accumulation of evidence in time. When subjects control stimulus duration, the decision terminates when the accumulated evidence reaches a criterion level. Under many natural circumstances and in many laboratory settings, the environment, rather than the subject, controls the stimulus duration. In these settings, it is generally assumed that subjects commit to a choice at the end of the stimulus stream. Indeed, failure to benefit from the full stream of information is interpreted as a sign of imperfect accumulation or memory leak. Contrary to these assumptions, we show that monkeys performing a direction discrimination task commit to a choice when the accumulated evidence reaches a threshold level (or bound), sometimes long before the end of stimulus. This bounded accumulation of evidence is reflected in the activity of neurons in the lateral intraparietal cortex. Thus, the readout of visual cortex embraces a termination rule to limit processing even when potentially useful information is available. 10.1523/JNEUROSCI.4761-07.2008

Dynamics of orientation tuning in macaque primary visual cortex.

DL Ringach — 2007-12-20T22:14:28-00:00

Nature, Vol. 387, No. 6630. (15 May 1997), pp. 281-284.

Orientation tuning of neurons is one of the chief emergent characteristics of the primary visual cortex, V1. Neurons of the lateral geniculate nucleus, which comprise the thalamic input to V1, are not orientation-tuned, but the majority of V1 neurons are quite selective. How orientation tuning arises within V1 is still controversial. To study this problem, we measured how the orientation tuning of neurons evolves with time using a new method: reverse correlation in the orientation domain. Orientation tuning develops after a delay of 30-45 milliseconds and persists for 40-85 ms. Neurons in layers 4C alpha or 4C beta, which receive direct input from the thalamus, show a single orientation preference which remains unchanged throughout the response period. In contrast, the preferred orientations of output layer neurons (in layers 2, 3, 4B, 5 or 6) usually change with time, and in many cases the orientation tuning may have more than one peak. This difference in dynamics is accompanied by a change in the sharpness of orientation tuning; cells in the input layers are more broadly tuned than cells in the output layers. Many of these observed properties of output layer neurons cannot be explained by simple feedforward models, whereas they arise naturally in feedback networks. Our results indicate that V1 is more than a bank of static oriented filters; the dynamics of output layer cells appear to be shaped by intracortical feedback.

A cellular mechanism of reward-related learning.

JN Reynolds — 2005-02-08T05:10:43-00:00

Nature, Vol. 413, No. 6851. (6 September 2001), pp. 67-70.

Positive reinforcement helps to control the acquisition of learned behaviours. Here we report a cellular mechanism in the brain that may underlie the behavioural effects of positive reinforcement. We used intracranial self-stimulation (ICSS) as a model of reinforcement learning, in which each rat learns to press a lever that applies reinforcing electrical stimulation to its own substantia nigra. The outputs from neurons of the substantia nigra terminate on neurons in the striatum in close proximity to inputs from the cerebral cortex on the same striatal neurons. We measured the effect of substantia nigra stimulation on these inputs from the cortex to striatal neurons and also on how quickly the rats learned to press the lever. We found that stimulation of the substantia nigra (with the optimal parameters for lever-pressing behaviour) induced potentiation of synapses between the cortex and the striatum, which required activation of dopamine receptors. The degree of potentiation within ten minutes of the ICSS trains was correlated with the time taken by the rats to learn ICSS behaviour. We propose that stimulation of the substantia nigra when the lever is pressed induces a similar potentiation of cortical inputs to the striatum, positively reinforcing the learning of the behaviour by the rats.

Suppression without inhibition in visual cortex.

TC Freeman — 2007-12-18T20:04:59-00:00

Neuron, Vol. 35, No. 4. (15 August 2002), pp. 759-771.

Neurons in primary visual cortex (V1) are thought to receive inhibition from other V1 neurons selective for a variety of orientations. Evidence for this inhibition is commonly found in cross-orientation suppression: responses of a V1 neuron to optimally oriented bars are suppressed by superimposed mask bars of different orientation. We show, however, that suppression is unlikely to result from intracortical inhibition. First, suppression can be obtained with masks drifting too rapidly to elicit much of a response in cortex. Second, suppression is immune to hyperpolarization (through visual adaptation) of cortical neurons responding to the mask. Signals mediating suppression might originate in thalamus, rather than in cortex. Thalamic neurons exhibit some suppression; additional suppression might arise from depression at thalamocortical synapses. The mechanisms of suppression are subcortical and possibly include the very first synapse into cortex.

Role of inhibition in the specification of orientation selectivity of cells in the cat striate cortex.

AB Bonds — 2007-12-17T21:23:17-00:00

Vis Neurosci, Vol. 2, No. 1. (1989), pp. 41-55.

Mechanisms supporting orientation selectivity of cat striate cortical cells were studied by stimulation with two superimposed sine-wave gratings of different orientations. One grating (base) generated a discharge of known amplitude which could be modified by the second grating (mask). Masks presented at nonoptimal orientations usually reduced the base-generated response, but the degree of reduction varied widely between cells. Cells with narrow orientation tuning tended to be more susceptible to mask presence than broadly tuned cells; similarly, simple cells generally showed more response reduction than did complex cells. The base and mask stimuli were drifted at different temporal frequencies which, in simple cells, permitted the identification of individual response components from each stimulus. This revealed that the reduction of the base response by the mask usually did not vary regularly with mask orientation, although response facilitation from the mask was orientation selective. In some sharply tuned simple cells, response reduction had clear local maxima near the limits of the cell's orientation-tuning function. Response reduction resulted from a nearly pure rightward shift of the response versus log contrast function. The lowest mask contrast yielding reduction was within 0.1-0.3 log unit of the lowest contrast effective for excitation. The temporal-frequency bandpass of the response-reduction mechanism resembled that of most cortical cells. The spatial-frequency bandpass was much broader than is typical for single cortical cells, spanning essentially the entire visual range of the cat. These findings are compatible with a model in which weak intrinsic orientation-selective excitation is enhanced in two stages: (1) control of threshold by nonorientation-selective inhibition that is continuously dependent on stimulus contrast; and (2) in the more narrowly tuned cells, orientation-selective inhibition that has local maxima serving to increase the slope of the orientation-tuning function.

Differential Attention-Dependent Response Modulation across Cell Classes in Macaque Visual Area V4.

JF Mitchell — 2007-07-09T14:47:29-00:00

Neuron, Vol. 55, No. 1. (5 July 2007), pp. 131-141.

The cortex contains multiple cell types, but studies of attention have not distinguished between them, limiting understanding of the local circuits that transform attentional feedback into improved visual processing. Parvalbumin-expressing inhibitory interneurons can be distinguished from pyramidal neurons based on their briefer action potential durations. We recorded neurons in area V4 as monkeys performed an attention-demanding task. We find that the distribution of action potential durations is strongly bimodal. Neurons with narrow action potentials have higher firing rates and larger attention-dependent increases in absolute firing rate than neurons with broad action potentials. The percentage increase in response is similar across the two classes. We also find evidence that attention increases the reliability of the neuronal response. This modulation is more than two-fold stronger among putative interneurons. These findings lead to the surprising conclusion that the strongest attentional modulation occurs among local interneurons that do not transmit signals between areas.

Spatial attention and the latency of neuronal responses in macaque area V4.

J Lee — 2007-10-03T03:24:26-00:00

J Neurosci, Vol. 27, No. 36. (5 September 2007), pp. 9632-9637.

The effects of attention on neuronal responses in visual cortex have been likened to a change in stimulus contrast. Attention and stimulus contrast both modulate the magnitude of neuronal responses. However, changes in stimulus contrast also affect the latency of visual responses. Although many neurophysiological studies have examined how attention affects the strength of neuronal responses, few have considered whether attention affects neuronal latencies. To compare directly the effects of stimulus contrast and attention, we recorded responses from individual neurons in area V4 of macaque monkeys while they performed a task that independently controlled spatial attention and stimulus contrast. As expected, changes in stimulus contrast affected both the magnitude and latency of neuronal responses. Although attention had the expected effects on the magnitudes of neuronal responses, we did not detect statistically reliable changes in neuronal latency. A direct comparison of the effects of contrast and attention revealed a reliable difference. When a shift in spatial attention decreased response magnitude, response latency increased much less than when the same magnitude change was caused by reducing stimulus contrast. Thus, attention is distinct from contrast in the way it affects the relationship between neuronal response magnitude and latency.

Do We Know What the Early Visual System Does?

Matteo Carandini — 2005-12-07T18:11:14-00:00

J. Neurosci., Vol. 25, No. 46. (16 November 2005), pp. 10577-10597.

We can claim that we know what the visual system does once we can predict neural responses to arbitrary stimuli, including those seen in nature. In the early visual system, models based on one or more linear receptive fields hold promise to achieve this goal as long as the models include nonlinear mechanisms that control responsiveness, based on stimulus context and history, and take into account the nonlinearity of spike generation. These linear and nonlinear mechanisms might be the only essential determinants of the response, or alternatively, there may be additional fundamental determinants yet to be identified. Research is progressing with the goals of defining a single "standard model" for each stage of the visual pathway and testing the predictive power of these models on the responses to movies of natural scenes. These predictive models represent, at a given stage of the visual pathway, a compact description of visual computation. They would be an invaluable guide for understanding the underlying biophysical and anatomical mechanisms and relating neural responses to visual perception.

Integration of visuospatial and effector information during symbolically cued limb movements in monkey lateral intraparietal area.

J Oristaglio — 2007-04-09T16:23:50-00:00

J Neurosci, Vol. 26, No. 32. (9 August 2006), pp. 8310-8319.

Natural behavior requires close but flexible coordination between attention, defined as selection for perception, and action. In recent years a distributed network including the lateral intraparietal area (LIP) has been implicated in visuospatial selection for attention and rapid eye movements (saccades), but the relation between the attentional and motor functions of this area remains unclear. Here we tested LIP neurons in a task that involved not an ocular but a manual operant response. Monkeys viewed a display containing one cue and several distractors and reported the orientation of the cue (right- or left-facing) by releasing one of two bars grasped, respectively, with the right or left hand. The movement in this task thus was associated with (cued by), but not directed toward, the visual stimulus. A large majority of neurons responded more when the cue rather than when a distractor was in their receptive field, suggesting that they contribute to the attentional selection of the cue. A fraction of these neurons also was modulated by limb release, thus simultaneously encoding cue location and the active limb. The results suggest that the LIP links behaviorally relevant visual information with motor variables relevant for solving a task in a wide range of circumstances involving goal-directed or symbolically cued movements and eye as well as limb movements. A central function of the LIP may be to coordinate visual and motor selection during a wide variety of behaviors.

Lateral habenula as a source of negative reward signals in dopamine neurons

Masayuki Matsumoto — 2007-05-24T01:27:28-00:00

Nature (23 May 2007)

Statistics of Midbrain Dopamine Neuron Spike Trains in the Awake Primate

Hannah Bayer — 2007-07-19T11:11:14-00:00

J Neurophysiol (5 July 2007), 01140.2006.

Work in behaving primates indicates that midbrain dopamine neurons encode a prediction error, the difference between an obtained reward and the reward expected. Studies of dopamine action potential timing in the alert and anaesthetized rat indicate that dopamine neurons respond in tonic and phasic modes, a distinction that has been less well characterized in the primates. We used spike train models to examine the relationship between the tonic and burst modes of activity in dopamine neurons while monkeys were performing a reinforced visuo-saccadic movement task. We studied spiking activity during four task-related intervals; two of these were intervals during which no task-related events occurred, while two were periods marked by task-related phasic activity. We found that dopamine neuron spike trains during the intervals when no events occurred were well described as tonic. Action potentials appeared to be independent, to occur at low frequency, and to be almost equally well described by Gaussian and Poisson-like (Gamma) processes. Unlike in the rat, interspike intervals as low as 20 ms were often observed during these presumptively tonic epochs. Having identified these periods of presumptively tonic activity we were able to quantitatively define phasic modulations (both increases and decreases in activity) during the intervals in which task-related events occurred. This analysis revealed that the phasic modulations of these neurons include both bursting, as has been described previously, and pausing. Together bursts and pauses seemed to provide a continuous, although non-linear, representation of the theoretically defined reward prediction error of reinforcement learning. 10.1152/jn.01140.2006

Monotonic coding of numerosity in macaque lateral intraparietal area.

JD Roitman — 2007-08-21T16:20:34-00:00

PLoS Biol, Vol. 5, No. 8. (August 2007)

As any child knows, the first step in counting is summing up individual elements, yet the brain mechanisms responsible for this process remain obscure. Here we show, for the first time, that a population of neurons in the lateral intraparietal area of monkeys encodes the total number of elements within their classical receptive fields in a graded fashion, across a wide range of numerical values (2-32). Moreover, modulation of neuronal activity by visual quantity developed rapidly, within 100 ms of stimulus onset, and was independent of attention, reward expectations, or stimulus attributes such as size, density, or color. The responses of these neurons resemble the outputs of "accumulator neurons" postulated in computational models of number processing. Numerical accumulator neurons may provide inputs to neurons encoding specific cardinal values, such as "4," that have been described in previous work. Our findings may explain the frequent association of visuospatial and numerical deficits following damage to parietal cortex in humans.

Behavioural improvements with thalamic stimulation after severe traumatic brain injury

ND Schiff — 2007-08-01T20:22:16-00:00

Nature, Vol. 448, No. 7153. (2007), pp. 600-603.

Probabilistic reasoning by neurons

Tianming Yang — 2007-06-04T03:11:35-00:00

Nature (03 June 2007)

Temporal Patterning of Saccadic Eye Movement Signals

Daniel Kimmel — 2007-07-19T13:42:35-00:00

J. Neurosci., Vol. 27, No. 29. (18 July 2007), pp. 7619-7630.

Electrical microstimulation is used widely in experimental neurophysiology to examine causal links between specific brain areas and their behavioral functions and is used clinically to treat neurological and psychiatric disorders in patients. Typically, microstimulation is applied to local brain regions as a train of equally spaced current pulses. We were interested in the sensitivity of a neural circuit to a train of variably spaced pulses, as is observed in physiological spike trains. We compared the effect of fixed, decelerating, accelerating, and randomly varying microstimulation patterns on the likelihood and metrics of eye movements evoked from the frontal eye field of monkeys, while holding the mean interpulse interval constant. Our results demonstrate that the pattern of microstimulation pulses strongly influences the probability of evoking a saccade, as well as the metrics of the saccades themselves. Specifically, the pattern most closely resembling physiological spike trains (accelerating pattern) was most effective at evoking a saccade, three times more so than the least effective decelerating pattern. A saccade-triggered average of effective random trains confirmed the positive relationship between accelerating rate and efficacy. These results have important implications for the use of electrical microstimulation in both experimental and clinical settings and suggest a means to study the role of temporal pattern in the encoding of behavioral and cognitive functions. 10.1523/JNEUROSCI.0386-07.2007

Lateral Habenula Stimulation Inhibits Rat Midbrain Dopamine Neurons through a GABAA Receptor-Mediated Mechanism

Huifang Ji — 2007-06-28T09:44:04-00:00

J. Neurosci., Vol. 27, No. 26. (27 June 2007), pp. 6923-6930.

Transient changes in the activity of midbrain dopamine neurons encode an error signal that contributes to associative learning. Although considerable attention has been devoted to the mechanisms contributing to phasic increases in dopamine activity, less is known about the origin of the transient cessation in firing accompanying the unexpected loss of a predicted reward. Recent studies suggesting that the lateral habenula (LHb) may contribute to this type of signaling in humans prompted us to evaluate the effects of LHb stimulation on the activity of dopamine and non-dopamine neurons of the anesthetized rat. Single-pulse stimulation of the LHb (0.5 mA, 100 micros) transiently suppressed the activity of 97% of the dopamine neurons recorded in the substantia nigra and ventral tegmental area. The duration of the cessation averaged [~]85 ms and did not differ between the two regions. Identical stimuli transiently excited 52% of the non-dopamine neurons in the ventral midbrain. Electrolytic lesions of the fasciculus retroflexus blocked the effects of LHb stimulation on dopamine neurons. Local application of bicuculline but not the SK-channel blocker apamin attenuated the effects of LHb stimulation on dopamine cells, indicating that the response is mediated by GABAA receptors. These data suggest that LHb-induced suppression of dopamine cell activity is mediated indirectly by orthodromic activation of putative GABAergic neurons in the ventral midbrain. The habenulomesencephalic pathway, which is capable of transiently suppressing the activity of dopamine neurons at a population level, may represent an important component of the circuitry involved in encoding reward expectancy. 10.1523/JNEUROSCI.0958-07.2007

The Neural Basis of Decision Making.

Joshua I Gold — 2007-07-02T13:48:33-00:00

Annu Rev Neurosci, Vol. 30 (21 July 2007), pp. 535-574.

The study of decision making spans such varied fields as neuroscience, psychology, economics, statistics, political science, and computer science. Despite this diversity of applications, most decisions share common elements including deliberation and commitment. Here we evaluate recent progress in understanding how these basic elements of decision formation are implemented in the brain. We focus on simple decisions that can be studied in the laboratory but emphasize general principles likely to extend to other settings.

Brain Reward Circuitry: Insights from Unsensed Incentives

Roy Wise — 2007-07-02T13:42:01-00:00

Neuron, Vol. 36, No. 2. (10 October 2002), pp. 229-240.

The natural incentives that shape behavior reach the central circuitry of motivation trans-synaptically, via the five senses, whereas the laboratory rewards of intracranial stimulation or drug injections activate reward circuitry directly, bypassing peripheral sensory pathways. The unsensed incentives of brain stimulation and intracranial drug injections thus give us tools to identify reward circuit elements within the associational portions of the CNS. Such studies have implicated the mesolimbic dopamine system and several of its afferents and efferents in motivational function. Comparisons of natural and laboratory incentives suggest hypotheses as to why some habits become compulsive and give insights into the roles of reinforcement and of prediction of reinforcement in habit formation.

What electrical microstimulation has revealed about the neural basis of cognition.

MR Cohen — 2007-06-11T17:10:47-00:00

Curr Opin Neurobiol, Vol. 14, No. 2. (April 2004), pp. 169-177.

Neurophysiologists have shown repeatedly that neural activity in different brain structures can be correlated with specific perceptual and cognitive functions, but the causal efficacy of the observed activity has generally been a matter of conjecture. By contrast, electrical microstimulation, which allows the experimenter to manipulate the activity of small groups of neurons with spatial and temporal precision, can now be used to demonstrate causal links between neural activity and specific cognitive functions. Here, we review this growing literature, including applications to the study of attention, visual and somatosensory perception, 'read-out' mechanisms for interpreting sensory maps, and contextual effects on perception. We also discuss potential applications of microstimulation to studies of higher cognitive functions such as decision-making and subjective experience.

Direct and indirect activation of cortical neurons by electrical microstimulation.

EJ Tehovnik — 2007-06-05T22:02:38-00:00

J Neurophysiol, Vol. 96, No. 2. (August 2006), pp. 512-521.

Electrical microstimulation has been used to elucidate cortical function. This review discusses neuronal excitability and effective current spread estimated by using three different methods: 1) single-cell recording, 2) behavioral methods, and 3) functional magnetic resonance imaging (fMRI). The excitability properties of the stimulated elements in neocortex obtained using these methods were found to be comparable. These properties suggested that microstimulation activates the most excitable elements in cortex, that is, by and large the fibers of the pyramidal cells. Effective current spread within neocortex was found to be greater when measured with fMRI compared with measures based on single-cell recording or behavioral methods. The spread of activity based on behavioral methods is in close agreement with the spread based on the direct activation of neurons (as opposed to those activated synaptically). We argue that the greater activation with imaging is attributed to transynaptic spread, which includes subthreshold activation of sites connected to the site of stimulation. The definition of effective current spread therefore depends on the neural event being measured.

Behavioral detection of electrical microstimulation in different cortical visual areas.

DK Murphey — 2007-05-22T14:23:42-00:00

Curr Biol, Vol. 17, No. 10. (15 May 2007), pp. 862-867.

The extent to which areas in the visual cerebral cortex differ in their ability to support perceptions has been the subject of considerable speculation. Experiments examining the activity of individual neurons have suggested that activity in later stages of the visual cortex is more closely linked to perception than that in earlier stages [1-9]. In contrast, results from functional imaging, transcranial magnetic stimulation, and lesion studies have been interpreted as showing that earlier stages are more closely coupled to perception [10-15]. We examined whether neuronal activity in early and later stages differs in its ability to support detectable signals by measuring behavioral thresholds for detecting electrical microstimulation in different cortical areas in two monkeys. By training the animals to perform a two-alternative temporal forced-choice task, we obtained criterion-free thresholds from five visual areas-V1, V2, V3A, MT, and the inferotemporal cortex. Every site tested yielded a reliable threshold. Thresholds varied little within and between visual areas, rising gradually from early to later stages. We similarly found no systematic differences in the slopes of the psychometric detection functions from different areas. These results suggest that neuronal signals of similar magnitude evoked in any part of visual cortex can generate percepts.

Neuronal switching of sensorimotor transformations for antisaccades.

M Zhang — 2007-05-10T21:48:07-00:00

Nature, Vol. 408, No. 6815. (8 2000), pp. 971-975.

The influence of cognitive context on orienting behaviour can be explored using the mixed memory-prosaccade, memory-antisaccade task. A symbolic cue, such as the colour of a visual stimulus, instructs the subject to make a brief, rapid eye movement (a saccade) either towards the stimulus (prosaccade) or in the opposite direction (antisaccade). Thus, the appropriate sensorimotor transformation must be switched on to execute the instructed task. Despite advances in our understanding of the neuronal processing of antisaccades, it remains unclear how the brain selects and computes the sensorimotor transformation leading to an antisaccade. Here we show that area LIP of the posterior parietal cortex is involved in these processes. LIP's population activity turns from the visual direction to the motor direction during memory-antisaccade trials. About one-third of the visual neurons in LIP produce a brisk, transient discharge in certain memory-antisaccade trials. We call this discharge 'paradoxical' because its timing is visual-like but its direction is motor. The paradoxical discharge shows, first, that switching occurs already at the level of visual cells, as previously proposed by Schlag-Rey and colleagues; and second, that this switching is accomplished very rapidly, within 50 ms from the arrival of the visual signals in LIP.

Progression in neuronal processing for saccadic eye movements from parietal cortex area lip to superior colliculus.

M Paré — 2007-05-07T16:13:08-00:00

J Neurophysiol, Vol. 85, No. 6. (June 2001), pp. 2545-2562.

Neurons in both the lateral intraparietal area (LIP) of the monkey parietal cortex and the intermediate layers of the superior colliculus (SC) are activated well in advance of the initiation of saccadic eye movements. To determine whether there is a progression in the covert processing for saccades from area LIP to SC, we systematically compared the discharge properties of LIP output neurons identified by antidromic activation with those of SC neurons collected from the same monkeys. First, we compared activity patterns during a delayed saccade task and found that LIP and SC neurons showed an extensive overlap in their responses to visual stimuli and in their sustained activity during the delay period. The saccade activity of LIP neurons was, however, remarkably weaker than that of SC neurons and never occurred without any preceding delay activity. Second, we assessed the dependence of LIP and SC activity on the presence of a visual stimulus by contrasting their activity in delayed saccade trials in which the presentation of the visual stimulus was either sustained (visual trials) or brief (memory trials). Both the delay and the presaccadic activity levels of the LIP neuronal sample significantly depended on the sustained presence of the visual stimulus, whereas those of the SC neuronal sample did not. Third, we examined how the LIP and SC delay activity relates to the future production of a saccade using a delayed GO/NOGO saccade task, in which a change in color of the fixation stimulus instructed the monkey either to make a saccade to a peripheral visual stimulus or to withhold its response and maintain fixation. The average delay activity of both LIP and SC neuronal samples significantly increased by the advance instruction to make a saccade, but LIP neurons were significantly less dependent on the response instruction than SC neurons, and only a minority of LIP neurons was significantly modulated. Thus despite some overlap in their discharge properties, the neurons in the SC intermediate layers showed a greater independence from sustained visual stimulation and a tighter relationship to the production of an impending saccade than the LIP neurons supplying inputs to the SC. Rather than representing the transmission of one processing stage in parietal cortex area LIP to a subsequent processing stage in SC, the differences in neuronal activity that we observed suggest instead a progressive evolution in the neuronal processing for saccades.

Top-Down Versus Bottom-Up Control of Attention in the Prefrontal and Posterior Parietal Cortices

Timothy Buschman — 2007-03-30T13:28:45-00:00

Science, Vol. 315, No. 5820. (30 March 2007), pp. 1860-1862.

Attention can be focused volitionally by "top-down" signals derived from task demands and automatically by "bottom-up" signals from salient stimuli. The frontal and parietal cortices are involved, but their neural activity has not been directly compared. Therefore, we recorded from them simultaneously in monkeys. Prefrontal neurons reflected the target location first during top-down attention, whereas parietal neurons signaled it earlier during bottom-up attention. Synchrony between frontal and parietal areas was stronger in lower frequencies during top-down attention and in higher frequencies during bottom-up attention. This result indicates that top-down and bottom-up signals arise from the frontal and sensory cortex, respectively, and different modes of attention may emphasize synchrony at different frequencies. 10.1126/science.1138071

Turning on and off recurrent balanced cortical activity.

Y Shu — 2006-11-30T21:35:21-00:00

Nature, Vol. 423, No. 6937. (15 May 2003), pp. 288-293.

The vast majority of synaptic connections onto neurons in the cerebral cortex arise from other cortical neurons, both excitatory and inhibitory, forming local and distant 'recurrent' networks. Although this is a basic theme of cortical organization, its study has been limited largely to theoretical investigations, which predict that local recurrent networks show a proportionality or balance between recurrent excitation and inhibition, allowing the generation of stable periods of activity. This recurrent activity might underlie such diverse operations as short-term memory, the modulation of neuronal excitability with attention, and the generation of spontaneous activity during sleep. Here we show that local cortical circuits do indeed operate through a proportional balance of excitation and inhibition generated through local recurrent connections, and that the operation of such circuits can generate self-sustaining activity that can be turned on and off by synaptic inputs. These results confirm the long-hypothesized role of recurrent activity as a basic operation of the cerebral cortex.

Center-surround organization of auditory receptive fields in the owl.

EI Knudsen — 2006-11-21T01:02:45-00:00

Science, Vol. 202, No. 4369. (17 November 1978), pp. 778-780.

The spinal receptive fields of specialized auditory units in the midbrain of the barn owl (Tyto abla) contain two functionally antagonistic areas: an excitatory center and an inhibitory surround. The response of these units represents the balance of acoustic activation of the two areas, which in turn depends upon the location, intensity, and spectral content of the sound stimulus.

Neuronal responses in area 7a to multiple stimulus displays: II. responses are suppressed at the cued location.

C Constantinidis — 2006-11-20T15:41:03-00:00

Cereb Cortex, Vol. 11, No. 7. (July 2001), pp. 592-597.

Everyday visual scenes contain a variety of stimuli that vary in their significance. The companion paper demonstrates that neurons in the posterior parietal cortex (PPC) are capable of encoding the spatial locations of the salient stimulus in multiple stimulus scenes. The present experiment sought to address how neuronal responses to stimuli appearing in the receptive field are modulated after attention has been drawn to one of multiple stimuli in a visual scene. We recorded from area 7a of the PPC in monkeys trained to do a spatial version of a match-to-sample task. The results show that neuronal responses are greatly suppressed when stimuli appear at previously attended locations. No reduction in responsiveness is observed for locations where stimuli had previously appeared but did not draw attention. These results support the hypothesis that area 7a has a role in redirecting attention to stimuli appearing at novel, unattended locations.

Neuronal responses in area 7a to multiple-stimulus displays: I. neurons encode the location of the salient stimulus.

C Constantinidis — 2006-11-20T15:37:34-00:00

Cereb Cortex, Vol. 11, No. 7. (July 2001), pp. 581-591.

The primate posterior parietal cortex (PPC) plays an important role in representing and recalling spatial relationships and in the ability to orient visual attention. This is evidenced by the parietal activation observed in brain imaging experiments performed during visuo- spatial tasks, and by the contralateral neglect syndrome that often accompanies parietal lesions. Individual neurons in monkey parietal cortex respond vigorously to the appearance of single, behaviorally relevant stimuli, but little is known about how they respond to more complex visual displays. The current experiments addressed this issue by recording activity from single neurons in area 7a of the PPC in monkeys performing a spatial version of a match-to-sample task. The task required them to locate salient stimuli in multiple-stimulus displays and release a lever after a subsequent stimulus appeared at the same location. Neurons responded preferentially to the appearance of salient stimuli inside their receptive fields. The presence of multiple stimuli did not affect appreciably the spatial tuning of responses in the majority of neurons or the population code for the location of the salient stimulus. Responses to salient stimuli could be distinguished from background stimuli approximately 100 ms after the onset of the cue. These results suggest that area 7a neurons represent the location of the stimulus attracting the animal's attention and can provide the spatial information required for directing attention to a salient stimulus in a complex scene.

Dopamine neurons can represent context-dependent prediction error.

H Nakahara — 2006-10-09T14:04:22-00:00

Neuron, Vol. 41, No. 2. (22 January 2004), pp. 269-280.

Midbrain dopamine (DA) neurons are thought to encode reward prediction error. Reward prediction can be improved if any relevant context is taken into account. We found that monkey DA neurons can encode a context-dependent prediction error. In the first noncontextual task, a light stimulus was randomly followed by reward, with a fixed equal probability. The response of DA neurons was positively correlated with the number of preceding unrewarded trials and could be simulated by a conventional temporal difference (TD) model. In the second contextual task, a reward-indicating light stimulus was presented with the probability that, while fixed overall, was incremented as a function of the number of preceding unrewarded trials. The DA neuronal response then was negatively correlated with this number. This history effect corresponded to the prediction error based on the conditional probability of reward and could be simulated only by implementing the relevant context into the TD model.