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Rif induces dorsal filopodia but the signalling pathway responsible for this has not been identified. We show here that Rif interacts with the I-BAR family protein IRTKS via its I-BAR domain. Rif also interacts with Pinkbar in N1E-115 mouse neuroblastoma cells. IRTKS and Rif induce dorsal membrane ruffles and filopodia. Dominant negative Rif inhibits the formation of IRTKS-induced morphological structures and Rif activity is blocked in IRTKS KO cells. To further define the Rif-IRTKS signalling pathway, we identify Eps8 and WAVE2 as IRTKS interactors. We find that Eps8 regulates the size and number of dorsal filopodia and membrane ruffles downstream of Rif-IRTKS, while WAVE2 modulates dorsal membrane ruffling. Furthermore, our data suggests that Tir, a protein essential for enterohemorrhagic E.coli infection, may compete for Rif for interaction with the I-BAR domain of IRKS. Based on these evidences we propose a model in which Rho family GTPases use the I-BAR proteins, IRSp53, IRTKS and Pinkbar, as a central mechanism to modulate cell morphology.

Rho GTPase Rif signals through IRTKS, Eps8 and WAVE2 to generate dorsal membrane ruffles and filopodia.