Neural responses are typically characterized by computing the mean firing rate. Yet response variability can exist across trials. Many studies have examined the impact of a stimulus on the mean response, yet few have examined the impact on response variability. We measured neural variability in 13 extracellularly-recorded datasets and one intracellularly-recorded dataset from 7 areas spanning the four cortical lobes. In every case, stimulus onset caused a decline in neural variability. This occurred even when the stimulus produced little change in mean firing rate. The variability decline was observable in membrane potential recordings, in the spiking of individual neurons, and in correlated spiking variability measured with implanted 96-electrode arrays. The variability decline was observed for all stimuli tested, regardless of whether the animal was awake, behaving, or anaesthetized. This widespread variability decline suggests a rather general property of cortex: that its state is stabilized by an input.
Several decades of psychophysical and neurophysiological studies have established that visual signals are enhanced at the locus of attention. What remains a mystery is the mechanism that initiates biases in the strength of visual representations. Recent evidence argues that, during spatial attention, these biases reflect nascent saccadic eye movement commands. We examined the functional interaction of saccade preparation and visual coding by electrically stimulating sites within the frontal eye fields (FEF) and measuring its effect on the activity of neurons in extrastriate visual cortex. Here we show that visual responses in area V4 could be enhanced after brief stimulation of retinotopically corresponding sites within the FEF using currents below that needed to evoke saccades. The magnitude of the enhancement depended on the effectiveness of receptive field stimuli as well as on the presence of competing stimuli outside the receptive field. Stimulation of non-corresponding FEF representations could suppress V4 responses. The results suggest that the gain of visual signals is modified according to the strength of spatially corresponding eye movement commands.
A large population of neurons can in principle produce an astronomical number of distinct firing patterns. In cortex however, these patterns lie in a space of lower dimension1-4, as if individual neurons were “obedient members of a huge orchestra”5. Here we use recordings from the visual cortex of mouse and monkey to investigate the relationship between individual neurons and the population, and to establish the underlying circuit mechanisms. We show that neighbouring neurons can differ in their coupling to the overall firing of the population, ranging from strongly coupled “choristers” to weakly coupled “soloists”. Population coupling is largely independent of sensory preferences, and it is a fixed cellular attribute, invariant to stimulus conditions. Neurons with high population coupling are more strongly affected by non-sensory behavioural variables such as motor intention. Population coupling reflects a causal relationship, predicting a neuron’s response to optogenetically-driven increases in local activity. Moreover, population coupling indicates synaptic connectivity: a neuron’s population coupling, measured in vivo, predicted subsequent in vitro estimates of the number of synapses received from its neighbours. Finally, population coupling provides a compact summary of population activity: knowledge of the population couplings of N neurons predicts a substantial portion of their N2 pairwise correlations. Population coupling therefore represents a novel, simple measure that characterises each neuron’s relationship to a larger population, explaining seemingly complex network firing patterns in terms of basic circuit variables.
Electrical microstimulation was used to study primary motor and premotor cortex in monkeys. Each stimulation train was 500 ms in duration, approximating the time scale of normal reaching and grasping movements and the time scale of the neuronal activity that normally accompanies movement. This stimulation on a behaviorally relevant time scale evoked coordinated, complex postures that involved many joints. For example, stimulation of one site caused the mouth to open and also caused the hand to shape into a grip posture and move to the mouth. Stimulation of this site always drove the joints toward this final posture, regardless of the direction of movement required to reach the posture. Stimulation of other cortical sites evoked different postures. Postures that involved the arm were arranged across cortex to form a map of hand positions around the body. This stimulation-evoked map encompassed both primary motor and the adjacent premotor cortex. We suggest that these regions fit together into a single map of the workspace around the body.
Moore, Tirin and Mazyar Fallah. Microstimulation of the frontal eye field and its effects on covert spatial attention. J Neurophysiol 91: 152-162, 2004. First published September 17, 2003 10.1152/jn.00741.2002. Many studies have established that the strength of visual perception and the strength of visual representations within visual cortex vary according to the focus of covert spatial attention. While it is clear that attention can modulate visual signals, the source of this modulation remains unknown. We have examined the possibility that saccade related mechanisms provide a source of spatial attention by studying the effects of electrical microstimulation of the frontal eye fields (FEF) on spatial attention. Monkeys performed a task in which they had to detect luminance changes of a peripheral target while ignoring a flashing distracter. The target luminance change could be preceded by stimulation of the FEF at current levels below that which evoked saccadic eye movements. We found that when the target change was preceded by stimulation of FEF, the monkey could detect smaller changes in target luminance. The increased sensitivity to the target change only occurred when the target was placed in the part of the visual field represented by neurons at the stimulation site. The magnitude of improvement depended on the temporal asynchrony of the stimulation onset and the target event. No significant effect of stimulation was observed when long intervals (Ͼ300 ms) between stimulation and the target event were used, and the magnitude of the increased sensitivity decreased systematically with increasing asynchrony. At the shortest asynchrony, FEF stimulation temporally overlapped the target event and the magnitude of the improvement was comparable to that of removing the distracter from the task. These results demonstrate that transient, but potent improvements in the deployment of covert spatial attention can be obtained by microstimulation of FEF sites from which saccadic eye movements are also evoked. I N T R O D U C T I O NSpatial attention refers to the selective processing of stimuli at behaviorally relevant locations at the expense of stimuli at other locations. Saccadic eye movements are the primary way by which visual resources (i.e., the foveas) are brought to bear on relevant spatial locations and are thus key exemplars of spatial attentive behavior. Much has been made of the fact that objects can be selected for attention without actually being fixated (Sperling and Melchner 1978), i.e., selected covertly, but a wealth of evidence suggests that the mechanisms of overt and covert attention are similar, if not the same (Corbetta et al.
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