Humans’ ability to recognize objects is remarkably robust across a variety of views unless faces are presented upside-down. Whether this face inversion effect (FIE) results from qualitative (distinct mechanisms) or quantitative processing differences (a matter of degree within common mechanisms) between upright and inverted faces has been intensely debated. Studies have focused on preferential responses to faces in face-specific brain areas, although face recognition also involves nonpreferential responses in non-face-specific brain areas. By using dynamic causal modeling with Bayesian model selection, here we show that dissociable cortical pathways are responsible for qualitative and quantitative mechanisms in the FIE in the distributed network for face recognition. When faces were upright, the early visual cortex (VC) and occipital and fusiform face areas (OFA, FFA) suppressed couplings to the lateral occipital cortex (LO), a primary locus of object processing. In contrast, they did not inhibit the LO when faces were inverted but increased couplings to the intraparietal sulcus, which has been associated with visual working memory. Furthermore, we found that upright and inverted face processing together involved the face network consisting of the VC, OFA, FFA, and inferior frontal gyrus. Specifically, modulatory connectivity within the common pathways (VC-OFA), implicated in the parts-based processing of faces, strongly correlated with behavioral FIE performance. The orientation-dependent dynamic reorganization of effective connectivity indicates that the FIE is mediated by both qualitative and quantitative differences in upright and inverted face processing, helping to resolve a central debate over the mechanisms of the FIE.
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