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Background  Rho1 is a small GTPase of the Ras superfamily that serves as the central component in a highly conserved signaling pathway that regulates tissue morphogenesis during development in all animals. Since there is tremendous diversity in the upstream signals that can activate Rho1 as well as the effector molecules that carry out its functions, it is important to define relevant  Rho1 -interacting genes for each morphogenetic event regulated by this signaling pathway. Previous work from our lab and others has shown that Rho signaling is necessary for the morphogenesis of leg imaginal discs during metamorphosis in  Drosophila , although a comprehensive identification of  Rho1 -interacting genes has not been attempted for this process.    Methodology/Principal Findings  We characterized an amorphic allele of  Rho1  that displays a poorly penetrant dominant malformed leg phenotype and is capable of being strongly enhanced by  Rho1 -interacting heterozygous mutations. We then used this allele in a second-site noncomplementation screen with the Exelixis collection of molecularly defined deficiencies to identify  Rho1 -interacting genes necessary for leg morphogenesis. In a primary screen of 461 deficiencies collectively uncovering ∼50% of the  Drosophila  genome, we identified twelve intervals harboring  Rho1 -interacting genes. Through secondary screening we identified six  Rho1 -interacting genes including three that were previously identified ( RhoGEF2 ,  broad , and  stubbloid ), thereby validating the screen. In addition, we identified  Cdc42 ,  Rheb  and  Sc2  as novel  Rho1 -interacting genes involved in adult leg development.    Conclusions/Significance  This screen identified well-known and novel  Rho1 -interacting genes necessary for leg morphogenesis, thereby increasing our knowledge of this important signaling pathway. We additionally found that Rheb may have a unique function in leg morphogenesis that is independent of its regulation of Tor.

A Second-Site Noncomplementation Screen for Modifiers of Rho1 Signaling during Imaginal Disc Morphogenesis in Drosophila