Characterizing modulatory interactions between areas V1 and V2 in human cortex: A new treatment of functional MRI data

Reversible cooling experiments in monkey visual cortex have demonstrated that visually driven neuronal activity in V2 depends on feedforward projections from V1, whereas neuronal activity in V1 is modulated by feedback, or reentrant, projections from V2. We present evidence for a homologous asymmetry in reciprocal connections between V1 and V2 in human cortex using physiological measurements obtained with functional MRI. The analysis was based on a nonlinear model of effective connectivity that partitioned the influence that one region exerted over another into an obligatory effect (an effect that depended only on the input) and a modulatory effect (an effect that represented an interaction between input and activity intrinsic to the target region). Using estimates of the modulatory effect we tested two related hypotheses: (1) that V2 would be a major source of modulatory influences on V1; and (2) that the modulatory effects of V2 on V1 would be greater than those of V1 on V2. The first constitutes a hypothesis about the regional or topographic organization of (modulatory) effective connectivity and the second hypothesis directly addresses the functional asymmetry suggested by reversible cooling experiments. The results confirmed that the origins of feedback modulatory effects on V1 were regionally specific and most pronounced in V2. In contrast, feedforward modulatory influences on V1 on V2 were negligible. This apparent asymmetry between feedforward and feedback modulatory interactions was evident in both hemispheres and appears to be a fairly robust feature of nonlinear interactions between striate and extrastriate cortex. © 1995 Wiley‐Liss, Inc.