Visual stimuli at locations beyond the classical receptive field (the area of the visual field from which responses can be evoked) can have profound effects on the selectivity and magnitude of neuronal responses in the visual cortex, even when these stimuli do not themselves evoke responses. It is generally assumed that intrinsic anatomical connections underlie these modulatory effects. Understanding how cortical neuron responses are shaped by such modulatory effects is important for understanding the role of cortical organization in determining cortical function, and in how the visual areas of the brain process complex real- world scenes. In this study we will clarify how neuronal responses in visual cortex are shaped by the spatial distribution of visual stimuli, and we will elucidate the anatomical substrates that link different points on the cortex (and therefore different spatial locations). This will therefore permit a direct test of whether local cortical connections do in fact underlie these modulatory surround effects. 1) In electrophysiological and neuroanatomical experiments in adult monkeys and ferrets, we will test whether the spatial extent and topography of response suppression or enhancement from beyond the classical receptive field can be accounted for by the distribution of local horizontal or cortical feedback connections within primary visual cortex (V1) and V2. 2) In electrophysiological studies in adult monkeys and ferrets, we will explore the stimulus specificity of response modulation from beyond the classical receptive field in V1 and V2. We will focus on V1 as the gateway to other cortical visual areas, but will also examine V2 to illuminate how general the properties and function of these modulatory effects are to all cortical visual areas. 3) In developmental studies in ferrets, we will systematically compare, across a large range of postnatal ages, the development of the modulatory surround effects to that of the responses in the classical receptive field, and to the refinement of intrinsic and extrinsic connections in V1. This will allow us to compare the developmental refinement of the modulatory effects to classical receptive field properties, and to further test whether intrinsic connections can underlie surround modulation. Comparison of details of the selectivity and spatial extent of different anatomical circuits with similar details from physiological investigations will provide information critical to an understanding of which anatomical circuits underlie particular functions, and in how neuronal function and connectivity underlie visual performance. This is a fundamental question in understanding how the cerebral cortex works, both in vision and in higher cognitive function.
