Kay KN, Winawer J, Mezer A, Wandell BA. Compressive spatial summation in human visual cortex. J Neurophysiol 110: 481-494, 2013. First published April 24, 2013 doi:10.1152/jn.00105.2013.-Neurons within a small (a few cubic millimeters) region of visual cortex respond to stimuli within a restricted region of the visual field. Previous studies have characterized the population response of such neurons using a model that sums contrast linearly across the visual field. In this study, we tested linear spatial summation of population responses using blood oxygenation level-dependent (BOLD) functional MRI. We measured BOLD responses to a systematic set of contrast patterns and discovered systematic deviation from linearity: the data are more accurately explained by a model in which a compressive static nonlinearity is applied after linear spatial summation. We found that the nonlinearity is present in early visual areas (e.g., V1, V2) and grows more pronounced in relatively anterior extrastriate areas (e.g., LO-2, VO-2). We then analyzed the effect of compressive spatial summation in terms of changes in the position and size of a viewed object. Compressive spatial summation is consistent with tolerance to changes in position and size, an important characteristic of object representation. (Fig. 1). The validity of this assumption is important to examine, as it affects the accuracy of pRF estimates and may reveal insight into response properties at different stages of the visual map hierarchy. Assessments of linearity of spatial summation have been conducted in both electrophysiology and fMRI, but these have provided conflicting conclusions (e.g., Britten and Heuer 1999;Hansen et al. 2004;Kastner et al. 2001;Pihlaja et al. 2008). Thus the precise nature of spatial pooling, and how well the linear approximation describes physiological responses, remains unclear.In this study, we examine spatial summation using systematic measurements of blood oxygenation level-dependent (BOLD) fMRI responses in human visual cortex to a range of spatial contrast patterns. We uncover a small nonlinear effect (subadditive spatial summation) in primary visual cortex and find that the nonlinear effect is pronounced in extrastriate maps. To account for the effect, we develop a computational model in which a compressive static nonlinearity is applied after linear spatial summation; this model substantially improves cross-validation performance compared with a linear spatial summation model.