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Background  Programmed cell death is used to remove excess cells between ommatidia in the  Drosophila  pupal retina. This death is required to establish the crystalline, hexagonal packing of ommatidia that characterizes the adult fly eye. In previously described  echinus  mutants, interommatidial cell sorting, which precedes cell death, occurred relatively normally. Interommatidial cell death was partially suppressed, resulting in adult eyes that contained excess pigment cells, and in which ommatidia were mildly disordered. These results have suggested that  echinus  functions in the pupal retina primarily to promote interommatidial cell death.    Results  We generated a number of new  echinu s alleles, some likely null mutants. Analysis of these alleles provides evidence that  echinus  has roles in cell sorting as well as cell death.  echinus  encodes a protein with homology to ubiquitin-specific proteases. These proteins cleave ubiquitin-conjugated proteins at the ubiquitin C-terminus. The  echinus  locus encodes multiple splice forms, including two proteins that lack residues thought to be critical for deubiquitination activity. Surprisingly, ubiquitous expression in the eye of versions of Echinus that lack residues critical for ubiquitin specific protease activity, as well as a version predicted to be functional, rescue the  echinus  loss-of-function phenotype. Finally, genetic interactions were not detected between  echinus  loss and gain-of-function and a number of known apoptotic regulators. These include Notch, EGFR, the caspases Dronc, Drice, Dcp-1, Dream, the caspase activators, Rpr, Hid, and Grim, the caspase inhibitor DIAP1, and Lozenge or Klumpfuss.    Conclusion  The  echinus  locus encodes multiple splice forms of a protein with homology to ubiquitin-specific proteases, but protease activity is unlikely to be required for  echinus  function, at least when  echinus  is overexpressed. Characterization of likely  echinus  null alleles and genetic interactions suggests that  echinus  acts at a novel point(s) to regulate interommatidial cell sorting and/or cell death in the fly eye.

bmc developmental biology

echinus, required for interommatidial cell sorting and cell death in the Drosophila pupal retina, encodes a protein with homology to ubiquitin-specific proteases