title: Horizontal transfer of an adaptive chimeric photoreceptor from bryophytes to ferns.
abstract: Ferns are well known for their shade-dwelling habits. Their ability to thrive under low-light conditions has been linked to the evolution of a novel chimeric photoreceptor--neochrome--that fuses red-sensing phytochrome and blue-sensing phototropin modules into a single gene, thereby optimizing phototropic responses. Despite being implicated in facilitating the diversification of modern ferns, the origin of neochrome has remained a mystery. We present evidence for neochrome in hornworts (a bryophyte lineage) and demonstrate that ferns acquired neochrome from hornworts via horizontal gene transfer (HGT). Fern neochromes are nested within hornwort neochromes in our large-scale phylogenetic reconstructions of phototropin and phytochrome gene families. Divergence date estimates further support the HGT hypothesis, with fern and hornwort neochromes diverging 179 Mya, long after the split between the two plant lineages (at least 400 Mya). By analyzing the draft genome of the hornwort Anthoceros punctatus, we also discovered a previously unidentified phototropin gene that likely represents the ancestral lineage of the neochrome phototropin module. Thus, a neochrome originating in hornworts was transferred horizontally to ferns, where it may have played a significant role in the diversification of modern ferns.
authors.affiliation: Department of Biology, Duke University, Durham, NC 27708.
authors.forename: Fay-Wei
authors.initials: FW
authors.lastname: Li
authors.forename: Juan Carlos
authors.initials: JC
authors.lastname: Villarreal
authors.forename: Steven
authors.initials: S
authors.lastname: Kelly
authors.forename: Carl J
authors.initials: CJ
authors.lastname: Rothfels
authors.forename: Michael
authors.initials: M
authors.lastname: Melkonian
authors.forename: Eftychios
authors.initials: E
authors.lastname: Frangedakis
authors.forename: Markus
authors.initials: M
authors.lastname: Ruhsam
authors.forename: Erin M
authors.initials: EM
authors.lastname: Sigel
authors.forename: Joshua P
authors.initials: JP
authors.lastname: Der
authors.forename: Jarmila
authors.initials: J
authors.lastname: Pittermann
authors.forename: Dylan O
authors.initials: DO
authors.lastname: Burge
authors.forename: Lisa
authors.initials: L
authors.lastname: Pokorny
authors.forename: Anders
authors.initials: A
authors.lastname: Larsson
authors.forename: Tao
authors.initials: T
authors.lastname: Chen
authors.forename: Stina
authors.initials: S
authors.lastname: Weststrand
authors.forename: Philip
authors.initials: P
authors.lastname: Thomas
authors.forename: Eric
authors.initials: E
authors.lastname: Carpenter
authors.forename: Yong
authors.initials: Y
authors.lastname: Zhang
authors.forename: Zhijian
authors.initials: Z
authors.lastname: Tian
authors.forename: Li
authors.initials: L
authors.lastname: Chen
authors.forename: Zhixiang
authors.initials: Z
authors.lastname: Yan
authors.forename: Ying
authors.initials: Y
authors.lastname: Zhu
authors.forename: Xiao
authors.initials: X
authors.lastname: Sun
authors.forename: Jun
authors.initials: J
authors.lastname: Wang
authors.forename: Dennis W
authors.initials: DW
authors.lastname: Stevenson
authors.forename: Barbara J
authors.initials: BJ
authors.lastname: Crandall-Stotler
authors.forename: A Jonathan
authors.initials: AJ
authors.lastname: Shaw
authors.forename: Michael K
authors.initials: MK
authors.lastname: Deyholos
authors.forename: Douglas E
authors.initials: DE
authors.lastname: Soltis
authors.forename: Sean W
authors.initials: SW
authors.lastname: Graham
authors.forename: Michael D
authors.initials: MD
authors.lastname: Windham
authors.forename: Jane A
authors.initials: JA
authors.lastname: Langdale
authors.forename: Gane Ka-Shu
authors.initials: GK
authors.lastname: Wong
authors.forename: Sarah
authors.initials: S
authors.lastname: Mathews
authors.forename: Kathleen M
authors.initials: KM
authors.lastname: Pryer
mesh_terms: D020418:Algal Proteins / Q000235:genetics; D049936:Anthocerotophyta / Q000235:genetics; D001483:Base Sequence; D044002:Bryophyta / Q000235:genetics*; D018744:DNA, Plant / Q000235:genetics; D019143:Evolution, Molecular; D029624:Ferns / Q000235:genetics*; D022761:Gene Transfer, Horizontal; D017343:Genes, Plant; D008969:Molecular Sequence Data; D055519:Photoreceptors, Plant / Q000235:genetics*; D056949:Phototropins / Q000235:genetics; D010802:Phylogeny; D010834:Phytochrome / Q000235:genetics; D011993:Recombinant Fusion Proteins / Q000235:genetics; D059467:Transcriptome; D024341:Xanthophylls / Q000235:genetics
chemical_list: D020418:Algal Proteins; D018744:DNA, Plant; D055519:Photoreceptors, Plant; D056949:Phototropins; D011993:Recombinant Fusion Proteins; D024341:Xanthophylls; C492072:neochrome; D010834:Phytochrome
keywords: chloroplast movement; phototropism
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