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Abstract

How attention shapes the structure of population activity has attracted substantial interest over the past decades. Attention has traditionally been associated with an increase in firing rates, reflecting a change in the gain of the population. More recent studies also report a change in noise correlations, which is thought to reflect changes in functional connectivity. However, since the degree of attention can vary substantially from trial to trial even within one experimental condition, the measured correlations could actually reflect fluctuations in the attention-related feedback signal (gain) rather than feed-forward noise, as often assumed. To gain insights into this issue we analytically analyzed the standard model of spatial attention, where directing attention to the receptive field of a neuron increases its response gain. We assumed conditionally independent neurons (no noise correlations) and asked how uncontrolled fluctuations in attention affect the correlation structure. First, we found that this simple model of spatial attention explains the empirically measured correlation structure quite well. In addition to a positive average level of correlations, it predicts both an increase in correlations with firing rates, as observed in many studies, and a decrease in correlations with the difference of two neurons’ tuning functions — a structure generally referred to as limited range correlations. Second, we asked how fluctuations in attention would affect the accuracy of a population code, if treated as noise by a downstream readout. Based on previous theoretical results, it would be expected that they negatively affect readout accuracy because of the limited range correlations they induce. Surprisingly, we found that this is not the case: correlations due to random gain fluctuations do not affect readout accuracy because their major axis is orthogonal to changes in the stimulus orientation. Our results can be readily generalized to include feature-based attention. The model has very few free parameters and can potentially account for a large fraction of the experimentally observed spike count (co-)variance.