Effect of Attentive Fixation in Macaque Thalamus and Cortex

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Abstract
Attentional modulation of neuronal responsiveness is common in many areas of visual cortex. We examined whether attentional modulation in the visual thalamus was quantitatively similar to that in cortex. Identical procedures and apparatus were used to compare attentional modulation of single neurons in seven different areas of the visual system: the lateral geniculate, three visual subdivisions of the pulvinar [inferior, lateral, dorsomedial part of lateral pulvinar (Pdm)], and three areas of extrastriate cortex representing early, intermediate, and late stages of cortical processing (V2, V4/PM, area 7a). A simple fixation task controlled transitions among three attentive states. The animal waited for a fixation point to appear (ready state), fixated the point until it dimmed (fixation state), and then waited idly to begin the next trial (idle state). Attentional modulation was estimated by flashing an identical, irrelevant stimulus in a neuron's receptive field during each of the three states; the three responses defined a “response vector” whose deviation from the line of equal response in all three states (the main diagonal) indicated the character and magnitude of attentional modulation. Attentional modulation was present in all visual areas except the lateral geniculate, indicating that modulation was of central origin. Prevalence of modulation was modest (26%) in pulvinar, and increased from 21% in V2 to 43% in 7a. Modulation had a push-pull character (as many cells facilitated as suppressed) with respect to the fixation state in all areas except Pdm where all cells were suppressed during fixation. The absolute magnitude of attentional modulation, measured by the angle between response vector and main diagonal expressed as a percent of the maximum possible angle, differed among brain areas. Magnitude of modulation was modest in the pulvinar (19–26%), and increased from 22% in V2 to 41% in 7a. However, average trial-to-trial variability of response, measured by the coefficient of variation, also increased across brain areas so that its difference among areas accounted for more than 90% of the difference in modulation magnitude among areas. We also measured attentional modulation by the ratio of cell discharge due to attention divided by discharge variability. The resulting signal-to-noise ratio of attention was small and constant, 1.3 ± 10%, across all areas of pulvinar and cortex. We conclude that the pulvinar, but not the lateral geniculate, is as strongly affected by attentional state as any area of visual cortex we studied and that attentional modulation amplitude is closely tied to intrinsic variability of response.