Drosophila, Shifted. Shifted is required for high levels of long-range signaling in the
developing wing imaginal disc. Surprisingly, shifted encodes the only Drosophila ortholog of
the secreted vertebrate protein Wnt Inhibitory Factor-1 (WIF-1), whose known role is to bind
to extracellular Wnts and inhibit their activity. However, Shifted does not regulate Hedgehog
signaling by affecting Wingless or Wnt signaling. We show instead that Shifted is a secreted …