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MBE Advance Access published online on October 4, 2007
Molecular Biology and Evolution, doi:10.1093/molbev/msm200
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© The Author 2007. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oxfordjournals.org

Research Article

Cytochrome P450 1 Genes in Early Deuterostomes (Tunicates and Sea Urchins) and Vertebrates (Chicken and Frog): Origin and Diversification of the CYP1 Gene Family

Jared V. Goldstone1, Heather.M.H. Goldstone2, Ann M. Morrison1, Ann Tarrant1, Suzanne E. Kern1, Bruce R. Woodin1 and John J. Stegeman1

1 Department of Biology, Woods Hole Oceanographic Institution, Woods Hole, MA 02543
2 Josephine Bay Paul Center, Marine Biological Laboratory, Woods Hole, MA 02543

Corresponding authors: Jared Goldstone, John Stegeman, Department of Biology, Redfield 3-42 MS#32, Woods Hole Oceanographic Institution, Woods Hole, MA 02543, Email: jgoldstone{at}whoi.edu or jstegeman{at}whoi.edu, Telephone: 508-289-4823 (JG) or 508-289-2320 (JS), Fax: 508-457-2134

Received for publication March 30, 2007. Revision received August 31, 2007. Accepted for publication September 5, 2007.

Cytochrome P450 family 1 (CYP1) proteins are important in a large number of toxicological processes. CYP1A and CYP1B genes are well known in mammals, but the evolutionary history of the CYP1 family as a whole is obscure; that history may provide insight into endogenous functions of CYP1 enzymes. Here we identify CYP1-like genes in early deuterostomes (tunicates and echinoderms), and several new CYP1 genes in vertebrates (chicken, Gallus gallus, and frog, Xenopus tropicalis). Profile Hidden Markov Models (HMMs) generated from vertebrate CYP1A and CYP1B protein sequences were used to identify five potential CYP1 homologs in the tunicate Ciona intestinalis genome. The C. intestinalis genes were cloned and sequenced, confirming the predicted sequences. Orthologs of four of these genes were found in the C. savignyi genome. Bayesian phylogenetic analyses group the tunicate genes in the CYP1 family, provisionally in two new subfamilies, CYP1E and CYP1F, which fall in the CYP1A and CYP1B/1C clades. Bayesian and maximum likelihood analyses predict functional divergence between the tunicate and vertebrate CYP1s, and regions within CYP substrate recognition sites were found to differ significantly in position-specific substitution rates between tunicates and vertebrates. Subsequently, ten CYP1-like genes were found in the echinoderm Strongylocentrotus purpuratus (sea urchin) genome. Several of the tunicate and echinoderm CYP1-like genes are expressed during development. Canonical xenobiotic response elements are present in the upstream genomic sequences of most tunicate and sea urchin CYP1s, and both groups are predicted to possess an aryl hydrocarbon receptor (AHR), suggesting possible regulatory linkage of AHR and these CYPs. The CYP1 family has undergone multiple rounds of gene duplication followed by functional divergence, with at least one gene lost in mammals. This study provides new insight into the origin and evolution of CYP1 genes.

Key Words: cytochrome P450 • CYP1A • CYP1B • CYP1C • substrate recognition site • functional selection


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