Table 1.

Drosophila heparan sulfate (HS) biosynthesis mutants

Mutant (gene)Protein (mammalian homolog)FunctionReferences
sugarless (sgl) UDP-glucose dehydrogenaseGAG biosynthesis; effects on Wg, FGF, Dpp signalingBinari et al., 1997; Hacker et al., 1997; Haerry et al., 1997; Toyoda et al., 2000a
sulfateless (sfl) N-deacetylase/N-sulfotransferaseHS modification (mainly in sulfation); effects on Wg, Hh, and FGF signalingBaeg et al., 2001; Lin et al., 1999; Lin and Perrimon, 1999; Toyoda et al., 2000a
tout-velu (ttv) EXT1HS co-polymerase; effects on Hh signaling and movement, Wg distribution, Dpp signaling and distributionBellaiche et al., 1998; Bornemann et al., 2004; Gallet et al., 2003; Han et al., 2004a; Takei et al., 2004; The et al., 1999; Toyoda et al., 2000a; Toyoda et al., 2000b
sister of ttv (sotv) EXT2HS co-polymerase; effects on Hh signaling and movement, Wg distribution, Dpp signaling and distributionBornemann et al., 2004; Han et al., 2004a; Takei et al., 2004
brother of ttv (botv) EXTL3HS polymerase; effects on Hh signaling and movement, Wg signaling and distribution, Dpp signaling and distributionHan et al., 2004a; Kim et al., 2002; Takei et al., 2004
fringe connection (frc) UDP-sugar transporterTransfer of UDP-glucuronic acid, UDP-N-acetylglucosamine and possibly UDP-xylose from the cytoplasm into the lumen of the ER/Golgi; effects on Wg, Hh, FGF, and fringe-dependent Notch signalingGoto et al., 2001; Selva et al., 2001
slalom Adenosine 3′-phosphate 5′-phosphosulfate transporterTransporter for adenosine 3′-phosphate 5′-phosphosulfate from the cytosol into the Golgi lumen; effects on Wg, Hh, FGF and DV patterning mediated by PipeKamiyama et al., 2003; Luders et al., 2003
  • Dpp. Decapentaplegic; ER, endoplasmic reticulum; FGF, fibroblast growth factor; GAG, glycosaminoglycan; HS, heparan sulfate; Hh, hedgehog; UDP, uridine 5′-diphosphate; Wg, Wingless.