“…Subsequent evidence, however, tended to favour the merging and interaction of colour and achromatic (Ach) contrast into mixed, shape-processing pathways (Sincich & Horton, 2005;Economides et al, 2011). In V1, colour-luminance neurons, tuned to orientation and spatial frequency, potentially provide a common form-processing pathway, with only a minority of lowpass, nonoriented (isotropic) neurons exclusively sensitive to colour (Thorell et al, 1984;Lennie et al, 1990;Leventhal et al, 1995;Schluppeck & Engel, 2002;Friedman et al, 2003;Johnson et al, 2008;Shapley & Hawken, 2011;Li et al, 2015). In V4, neurophysiological evidence suggests multiple functional organizations, including colourselective neurons that process shape (Bushnell et al, 2011;Bushnell & Pasupathy, 2012), segregated pathways with colour contrast or luminance contrast preferences, and colour-luminance shape-processing regions (Tanigawa et al, 2010).…”