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Wyeth Bair

Publications and source records attributed to Wyeth Bair.

9 recordsLinked to original sources

Dynamics of suppression in macaque primary visual cortex.

The response of a neuron in primary visual cortex (V1) to an optimal stimulus in its classical receptive field (CRF) can be reduced by the presence of an orthogonal mask, a phenomenon known as cross-orientation suppression. The presence of a parallel stimulus outside the CRF can have a similar effect, in this case known as surround suppression. We used a novel stimulus to probe the time course of cross-orientation suppression and found that it is very fast, starting even before the response to optimal excitatory stimuli. However, it occurs with some delay after the offset response, considered to be a measure of the earliest excitatory signals that reach the CRF. We also examined the time course of response to a stimulus presented outside the CRF and found that cross-orientation suppression begins substantially earlier than surround suppression measured in the same cells. Together, these findings suggest that cross-orientation suppression is attributable to either direct feedforward signal paths to V1 neurons or a circuit involving fast local interneurons within V1. Feedback from higher cortical areas is implicated in surround suppression, but our results make this an implausible mechanism for cross-orientation suppression. We conclude that suppression from inside and outside the CRF occur through different mechanisms.

Action Potentials↗

Visual receptive field organization.

Increasingly systematic approaches to quantifying receptive fields in primary visual cortex, combined with inspired ideas about functional circuitry, non-linearities, and visual stimuli, are bringing new interest to classical problems. This includes the distinction and hierarchy between simple and complex cells, the mechanisms underlying the receptive field surround, and debates about optimal stimuli for mapping receptive fields. An important new problem arises from recent observations of stimulus-dependent spatial and temporal summation in primary visual cortex. It appears that the receptive field can no longer be considered unique, and we might have to relinquish this cherished notion as the embodiment of neuronal function in primary visual cortex.

Animals↗

Adaptive temporal integration of motion in direction-selective neurons in macaque visual cortex.

Direction-selective neurons in the primary visual cortex (V1) and the extrastriate motion area MT/V5 constitute a critical channel that links early cortical mechanisms of spatiotemporal integration to downstream signals that underlie motion perception. We studied how temporal integration in direction-selective cells depends on speed, spatial frequency (SF), and contrast using randomly moving sinusoidal gratings and spike-triggered average (STA) analysis. The window of temporal integration revealed by the STAs varied substantially with stimulus parameters, extending farther back in time for slow motion, high SF, and low contrast. At low speeds and high SF, STA peaks were larger, indicating that a single spike often conveyed more information about the stimulus under conditions in which the mean firing rate was very low. The observed trends were similar in V1 and MT and offer a physiological correlate for a large body of psychophysical data on temporal integration. We applied the same visual stimuli to a model of motion detection based on oriented linear filters (a motion energy model) that incorporated an integrate-and-fire mechanism and found that it did not account for the neuronal data. Our results show that cortical motion processing in V1 and in MT is highly nonlinear and stimulus dependent. They cast considerable doubt on the ability of simple oriented filter models to account for the output of direction-selective neurons in a general manner. Finally, they suggest that spike rate tuning functions may miss important aspects of the neural coding of motion for stimulus conditions that evoke low firing rates.

Action Potentials↗

Time course and time-distance relationships for surround suppression in macaque V1 neurons.

Iso-orientation surround suppression is a powerful form of visual contextual modulation in which a stimulus of the preferred orientation of a neuron placed outside the classical receptive field (CRF) of the neuron suppresses the response to stimuli within the CRF. This suppression is most often attributed to orientation-tuned signals that propagate laterally across the cortex, activating local inhibition. By studying the temporal properties of surround suppression, we have uncovered characteristics that challenge standard notions of surround suppression. We found that the latency of suppression depended on its strength. Across cells, strong suppression arrived on average 30 msec earlier than weak suppression, and suppression sometimes arrived faster than the excitatory CRF response. We compared the relative latency of CRF response onset and offset with the relative latency of suppression onset and offset. Response onset was delayed relative to response offset in the CRF but not in the surround. This is not the expected result if neurons targeted by suppression are like those that generate it. We examined the time course of suppression as a function of distance of the surround stimulus from the CRF and found that suppression was predominantly sustained for nearby stimuli and predominantly transient for distant stimuli. By comparing the latency of suppression for nearby and distant stimuli, we found that orientation-tuned suppression could effectively propagate across 6 - 8 mm of cortex at approximately 1 m/sec. This is considerably faster than expected for horizontal cortical connections previously implicated in surround suppression. We offer refinements to circuits for surround suppression that account for these results and describe how feedback from cells with large CRFs can account for the rapid propagation of suppression within V1.

Animals↗

Signals in macaque striate cortical neurons that support the perception of glass patterns.

Glass patterns are texture stimuli made by pairing randomly placed dots with partners at specific offsets. The strong percept of global form that arises from the sparse local orientation cues has made these patterns the subject of psychophysical investigations, yet neuronal responses to Glass patterns have not been studied. We measured the responses of neurons in macaque striate cortex (V1) to dynamic, translational Glass patterns as a function of dot separation and dot-pair orientation. Responses were selective, but were on average more than an order of magnitude weaker than responses to sinusoidal gratings. Response and selectivity were greatest when the dot-pair orientation matched that of the preferred grating and when dot separation was between one-quarter and one-half of the spatial period of the optimal grating; changing the dot-pair separation or inverting the contrast of one of the dots radically changed the orientation selectivity. We computed the expected responses for a receptive field model to translational Glass patterns and found that the complexity of our V1 tuning curves could be understood in terms of the responses of linear filters to pairs of dots. This modeling connects our understanding of V1 receptive fields as rectified, quasi-linear filters with results from psychophysical studies of Glass patterns. Our results provide a basis for studying how subsequent visual areas integrate weak, local signals into global form percepts.

Action Potentials↗

The timing of response onset and offset in macaque visual neurons.

We used fast, pseudorandom temporal sequences of preferred and antipreferred stimuli to drive neuronal firing rates rapidly between minimal and maximal across the visual system. Stimuli were tailored to the preferences of cells recorded in the lateral geniculate nucleus (magnocellular and parvocellular), primary visual cortex (simple and complex), and the extrastriate motion area MT. We found that cells took longer to turn on (to increase their firing rate) than to turn off (to reduce their rate). The latency difference (onset minus offset) varied from several to tens of milliseconds across cell type and stimulus class and was correlated with spontaneous or driven firing rates for most cell classes. The delay for response onset depended on the nature of the stimulus present before the preferred stimulus appeared, and may result from persistent inhibition caused by antipreferred stimuli or from suppression that followed the offset of the preferred stimulus. The onset delay showed three distinct types of dependence on the temporal sequence of stimuli across classes of cells, implying that suppression may accumulate or wear off with time. Onset latency is generally longer, can be more variable, and has marked stimulus dependence compared with offset latency. This suggests an important role for offset latency in assessing the speed of information transmission in the visual system and raises the possibility that signal offsets provide a timing reference for visual processing. We discuss the origin of the delay in onset latency compared with offset latency and consider how it may limit the utility of certain feedforward circuits.

Action Potentials↗

Nature and interaction of signals from the receptive field center and surround in macaque V1 neurons.

Information is integrated across the visual field to transform local features into a global percept. We now know that V1 neurons provide more spatial integration than originally thought due to the existence of their nonclassical inhibitory surrounds. To understand spatial integration in the visual cortex, we have studied the nature and extent of center and surround influences on neuronal response. We used drifting sinusoidal gratings in circular and annular apertures to estimate the sizes of the receptive field's excitatory center and suppressive surround. We used combinations of stimuli inside and outside the receptive field to explore the nature of the surround influence on the receptive field center as a function of the relative and absolute contrast of stimuli in the two regions. We conclude that the interaction is best explained as a divisive modulation of response gain by signals from the surround. We then develop a receptive field model based on the ratio of signals from Gaussian-shaped center and surround mechanisms. We show that this model can account well for the variations in receptive field size with contrast that we and others have observed and for variations in size with the state of contrast adaptation. The model achieves this success by simple variations in the relative gain of the two component mechanisms of the receptive field. This model thus offers a parsimonious explanation of a variety of phenomena involving changes in apparent receptive field size and accounts for these phenomena purely in terms of two receptive field mechanisms that do not themselves change in size. We used the extent of the center mechanism in our model as an indicator of the spatial extent of the central excitatory portion of the receptive field. We compared the extent of the center to measurements of horizontal connections within V1 and determined that horizontal intracortical connections are well matched in extent to the receptive field center mechanism. Input to the suppressive surround may come in part from feedback signals from higher areas.

Adaptation, Physiological↗

Selectivity and spatial distribution of signals from the receptive field surround in macaque V1 neurons.

The responsiveness of neurons in V1 is modulated by stimuli placed outside their classical receptive fields. This nonclassical surround provides input from a larger portion of the visual scene than originally thought, permitting integration of information at early levels in the visual processing stream. Signals from the surround have been reported variously to be suppressive and facilitatory, selective and unselective. We tested the specificity of influences from the surround by studying the interactions between drifting sinusoidal gratings carefully confined to conservatively defined center and surround regions. We found that the surround influence was always suppressive when the surround grating was at the neuron's preferred orientation. Suppression tended to be stronger when the surround grating also moved in the neuron's preferred direction, rather than its opposite. When the orientation in the surround was 90 degrees from the preferred orientation (orthogonal), suppression was weaker, and facilitation was sometimes evident. The tuning of surround signals therefore tended to match the tuning of the center, though the tuning of the surround was somewhat broader. The tuning of suppression also depended on the contrast of the center grating-when the center grating was reduced in contrast, orthogonal surround stimuli became relatively more suppressive. We also found evidence for the tuning of the surround being dependent to some degree on the stimulus used in the center-suppression was often stronger for a given center stimulus when the parameters of the surround grating matched the parameters of the center grating even when the center grating was not itself of the optimal direction or orientation. We also explored the spatial distribution of surround influence and found an orderly relationship between the orientation of grating patches presented to regions of the surround and the position of greatest suppression. When surround gratings were oriented parallel to the preferred orientation of the receptive field, suppression was strongest at the receptive field ends. When surround gratings were orthogonal, suppression was strongest on the flanks. We conclude that the surround has complex effects on responses from the classical receptive field. We suggest that the underlying mechanism of this complexity may involve interactions between relatively simple center and surround mechanisms.

Algorithms↗

No doubt about offset latency.

Neuronal response latency usually refers to the time between the presentation of a visual stimulus and the elevation in firing rate that follows. Expanding on this idea, the concept of response offset latency refers to the time between the removal of a stimulus (or its replacement with one that is less effective) and the resulting decline in firing rate. The initial observation that offset latency is usually shorter than onset latency (Bair et al., 2002) has been called into question on the basis of the pulsatile nature of visual stimuli presented on a CRT (Gawne & Woods, 2003). Here, a counter argument is presented in support of the results of Bair et al., 2002.

Animals↗