title: Genomic organization of duplicated short wave-sensitive and long wave-sensitive opsin genes in the green swordtail, Xiphophorus helleri.
abstract: BACKGROUND
  Long wave-sensitive (LWS) opsin genes have undergone multiple lineage-specific duplication events throughout the evolution of teleost fishes. LWS repertoire expansions in live-bearing fishes (family Poeciliidae) have equipped multiple species in this family with up to four LWS genes. Given that color vision, especially attraction to orange male coloration, is important to mate choice within poeciliids, LWS opsins have been proposed as candidate genes driving sexual selection in this family. To date the genomic organization of these genes has not been described in the family Poeciliidae, and little is known about the mechanisms regulating the expression of LWS opsins in any teleost.
  
  
  RESULTS
  Two BAC clones containing the complete genomic repertoire of LWS opsin genes in the green swordtail fish, Xiphophorus helleri, were identified and sequenced. Three of the four LWS loci identified here were linked in a tandem array downstream of two tightly linked short wave-sensitive 2 (SWS2) opsin genes. The fourth LWS opsin gene, containing only a single intron, was not linked to the other three and is the product of a retrotransposition event. Genomic and phylogenetic results demonstrate that the LWS genes described here share a common evolutionary origin with those previously characterized in other poeciliids. Using qualitative RT-PCR and MSP we showed that each of the LWS and SWS2 opsins, as well as three other cone opsin genes and a single rod opsin gene, were expressed in the eyes of adult female and male X. helleri, contributing to six separate classes of adult retinal cone and rod cells with average lambdamax values of 365 nm, 405 nm, 459 nm, 499 nm, 534 nm and 568 nm. Comparative genomic analysis identified two candidate teleost opsin regulatory regions containing putative CRX binding sites and hormone response elements in upstream sequences of LWS gene regions of seven teleost species, including X. helleri.
  
  
  CONCLUSIONS
  We report the first complete genomic description of LWS and SWS2 genes in poeciliids. These data will serve as a reference for future work seeking to understand the relationship between LWS opsin genomic organization, gene expression, gene family evolution, sexual selection and speciation in this fish family.
authors.affiliation: Department of Biological Sciences, Simon Fraser University, Burnaby, BC, Canada.
authors.forename: Corey T
authors.initials: CT
authors.lastname: Watson
authors.forename: Krzysztof P
authors.initials: KP
authors.lastname: Lubieniecki
authors.forename: Ellis
authors.initials: E
authors.lastname: Loew
authors.forename: William S
authors.initials: WS
authors.lastname: Davidson
authors.forename: Felix
authors.initials: F
authors.lastname: Breden
mesh_terms: D000818:Animals; D001483:Base Sequence; D022202:Chromosomes, Artificial, Bacterial; D003118:Color Perception / Q000235:genetics / Q000502:physiology; D003532:Cyprinodontiformes / Q000235:genetics*; D019143:Evolution, Molecular; D005260:Female; D029941:Fish Proteins / Q000235:genetics*; D015723:Gene Library; D020131:Genes, Duplicate; D008297:Male; D008969:Molecular Sequence Data; D010802:Phylogeny; D017299:Rod Opsins / Q000235:genetics*; D017422:Sequence Analysis, DNA; D026801:Synteny
chemical_list: D029941:Fish Proteins; D017299:Rod Opsins