[2] Helene George and Regine Terracol discovered the first vrille alleles (vri1 and vri2) in 1997 by EMS-mutagenesis assay and found their products to be transcription factors involved in embryogenesis.
[3] Justin Blau elucidated additional "vrille" implications in 1999 while screening for clock-controlled genes in Drosophila heads that responded to PER/TIM heterodimers.
[7] Vrille acts as an enhancer of decapentaplegic (dpp) and easter, genes critical to the development of dorsal/ventral axis in the process of regional differentiation during Drosophila embryogenesis.
Easter is involved in initiating a protease cascade that activates the dorsal gene, resulting in repression of dpp in the ventral portion of the embryo during early fly development.
Overexpression of vri causes anti-proliferative effects in processes vital for limb generation, as well as abnormal phenotypes in salivary glands and internal organs.
The crustacean Daphnia magna co-opts its vri ortholog for activating the doublesex sex-determination gene, a task accomplished by tra in flies.