Institute of Cognitive Neuroscience and Department ofPsychology, University College London, Alexandra House,17 Queen Square, London WC1N 3AR, UKSummaryFaces contain structural information, for identifying individ-uals, as well as changeable information, which can conveyemotion and direct attention. Neuroimaging studies revealbrain regions thatexhibit preferentialresponses toinvariant[1, 2] or changeable [3–5] facial aspects but the functionalconnections between these regions are unknown. We ad-dressed this issue by causally disrupting two face-selectiveregions with thetaburst transcranial magnetic stimulation(TBS) and measuring the effects of this disruption in localand remote face-selective regions with functional magneticresonance imaging (fMRI). Participants were scanned, overtwo sessions, while viewing dynamic or static faces and ob-jects. During these sessions, TBS was delivered over theright occipital face area (rOFA) or right posterior superiortemporal sulcus (rpSTS). Disruption of the rOFA reducedthe neural response to both static and dynamic faces inthe downstream face-selective region in the fusiform gyrus.In contrast, the response to dynamic and static faces wasdoubly dissociated in the rpSTS. Namely, disruption of therOFA reduced the response to static but not dynamic faces,while disruption of the rpSTS itself reduced the response todynamic but not static faces. These results suggest thatdynamic and static facial aspects are processed via dis-sociable cortical pathways that begin in early visual cortex,a conclusion inconsistent with current models of faceperception [6–9].ResultsInfluential models of face perception [6–8] propose that twofunctionally distinct cortical pathways process different facialaspects. The ventral pathway, which includes the fusiformface area (FFA) [10], preferentially responds to invariant facialaspects, such as individual identity. The lateral pathway,which includes the posterior superior temporal sulcus(pSTS) [3], preferentially responds to changeable facial as-pects, such as emotional expression and eye-gaze direction[4]. Crucially, despite functional differences, both pathwaysare believed to begin in the same face-selective region, theoccipital face area (OFA) [7–9, 11]. In the present study wecausally tested the hypothesis that the OFA is the solegateway for the face perception network using a ‘‘virtual’’lesion approach.Toexaminehowface-selectiveregionsarefunctionallycon-nected,thetabursttranscranialmagneticstimulation(TBS)[12]was used to transiently disrupt the brains of neurologicallyhealthy participants. The effects of this disruption were thenmeasuredinlocalandremoteface-selectiveregionswithfunc-tionalmagneticresonanceimaging (fMRI).Wereasonedthat ifthe OFA is the sole source of dynamic and static face informa-tion for the FFA and pSTS, then disrupting the OFA wouldreduce the neural response to dynamic and static faces inboth the FFA and pSTS. However, if a separate pathwayconveying only dynamic face information exists to the pSTS,independently of the OFA, then disruption of the OFA wouldhave relatively little impact on the response to dynamic facesin the pSTS (see Figure 1).Participants completed two scanning sessions, performedon separate days, while viewing face and object stimulithat were either dynamic or static (see Figure 2). Scanningwas performed before and after TBS was delivered over thefunctionally localized right OFA (rOFA) or right posterior supe-rior temporal sulcus (rpSTS). We then measured what ef-fect TBS disruption had on the neural response in both thestimulated regions (rOFA and rpSTS), as well as in the rightFFA (rFFA), a face-selective region on the ventral corticalsurface that cannot be directly stimulated by TBS. The magni-tuderesponsesfromeachface-selectiveROIaswellasfortheright extrastriate body area (rEBA) and right lateral occipitalarea (rLO) are shown in full in Figures S1 and S2 availableonline.ROI AnalysisTo understand what effect TBS stimulation had on the threeface-selective regions, we calculated the size of the TBSdisruptive effect in the rpSTS, rOFA, and rFFA. This wasdone by subtracting the BOLD responses for each stimuluscategory (dynamic faces, static faces, dynamic objects, staticobjects) after TBS stimulation of the rOFA and rpSTS from thepre-TMS baseline response in each ROI (see Figure 3).Thedatawerethenentered intoa2(TMS: TBStorOFA;TBSto rpSTS) by 2 (motion: dynamic, static) by 2 (stimulus: faces,objects) by 3 (ROI: rFFA, rpSTS; rOFA) repeated-measuresANOVA. Results showed a main effect of stimulus (F (1,14) =7.3, p = 0.017) as well as interactions between motion andTMS (F (1,14) = 4.2, p = 0.048) and between motion, stimulus,and TMS (F (1,14) = 3.6, p = 0.041). Crucially, there was alsoa significant interaction between ROI, motion, stimulus, andTMS(F(2,28)=3,p=0.043).Nootherinteractionsapproachedsignificance.Separate ANOVAs performed in each of the face-selectiveROIs (reported in full in Supplemental Information) demon-strated that TBS stimulation of the rOFA and rpSTS induceda double dissociation between the response to dynamic andstatic faces in the rpSTS (Figure 3). The response to staticfaces in the rpSTS was reduced by stimulation of the rOFAbut not of the rpSTS, while the response to dynamic faces inthe rpSTS was reduced by stimulation of the rpSTS but notof the rOFA. This result is consistent with the hypothesis thatdynamic face information can reach the rpSTS independently