Armadillo nuclear import is regulated by cytoplasmic anchor Axin and nuclear anchor dTCF/Pan

Drosophila melanogaster Armadillo plays two distinct roles during developme nt. It is a component of adherens junctions, and functions as a transcripti onal activator in response to Wingless signaling. In the current model, Win gless signal causes stabilization of cytoplasmic Armadillo allowing it to e nter the nucleus where it can activate transcription. However, the mechanis m of nuclear import and export remains to be elucidated. In this study, we show that two gain-of-function alleles of Armadillo activate Wingless signa ling by different mechanisms. The S10 allele was previously found to locali ze to the nucleus, where it activates transcription. In contrast, the Delta Arm allele localizes to the plasma membrane, and forces endogenous Arm int o the nucleus. Therefore, Delta Arm is dependent on the presence of a funct ional endogenous allele of arm to activate transcription. We show that Delt a Arm may function by titrating Axin protein to the membrane, suggesting th at it acts as a cytoplasmic anchor keeping Arm out of the nucleus. In axin mutants, Arm is localized to the nuclei, We find that nuclear retention is dependent on dTCF/Pangolin, This suggests that cellular distribution of Arm is controlled by an anchoring system, where various nuclear and cytoplasmi c binding partners determine its localization.