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UDP-glucuronosyltransferases (Ugts) are a supergene family of phase II drug-metabolizing enzymes that catalyze the conjugation of numerous hydrophobic small molecules with the UDP-glucuronic acid, converting them into hydrophilic molecules. Here, we report the identification and cloning of the complete zebrafish  Ugt  gene repertoire. We found that the zebrafish genome contains 45  Ugt  genes that can be divided into three families:  Ugt1 ,  Ugt2 , and  Ugt5 . Both  Ugt1  and  Ugt2  have two unlinked clusters:  a  and  b . The  Ugt1a ,  Ugt1b ,  Ugt2a , and  Ugt2b  clusters each contain variable and constant regions, similar to that of the protocadherin ( Pcdh ), immunoglobulin ( Ig ), and T-cell receptor ( Tcr ) clusters. Cloning the full-length coding sequences confirmed that each of the variable exons is separately spliced to the set of constant exons within each zebrafish  Ugt  cluster. Comparative analyses showed that both  a  and  b  clusters of the zebrafish  Ugt1  and  Ugt2  genes have orthologs in other teleosts, suggesting that they may be resulted from the “fish-specific” whole-genome duplication event. The  Ugt5  genes are a novel family of  Ugt  genes that exist in teleosts and amphibians. Their entire open reading frames are encoded by single large exons. The zebrafish  Ugt1 ,  Ugt2 , and  Ugt5  genes can generate additional transcript diversity through alternative splicing. Based on phylogenetic analyses, we propose that the ancestral tetrapod and teleost  Ugt1  clusters contained multiple  Ugt1  paralogs. After speciation, these ancestral  Ugt1  clusters underwent lineage-specific gene loss and duplication. The ancestral vertebrate  Ugt2  cluster also underwent lineage-specific duplication. The intronless  Ugt5  open reading frames may be derived from retrotransposition followed by gene duplication. They have been expanded dramatically in teleosts and have become the most abundant  Ugt  family in these lineages. These findings have interesting implications regarding the molecular evolution of genes with diversified variable exons in vertebrates.

Cloning and Comparative Analyses of the Zebrafish Ugt Repertoire Reveal Its Evolutionary Diversity