title: MAJOR DEVELOPMENTAL REGULATORS AND THEIR EXPRESSION IN TWO CLOSELY RELATED SPECIES OF PORPHYRA (RHODOPHYTA)(1).
abstract: Little is known about the genetic and biochemical mechanisms that underlie red algal development, for example, why the group failed to evolve complex parenchyma and tissue differentiation. Here we examined expressed sequence tag (EST) data from two closely related species, Porphyra umbilicalis (L.) J. Agardh and P. purpurea (Roth) C. Agardh, for conserved developmental regulators known from model eukaryotes, and their expression levels in several developmental stages. Genes for most major developmental families were present, including MADS-box and homeodomain (HD) proteins, SNF2 chromatin-remodelers, and proteins involved in sRNA biogenesis. Some of these genes displayed altered expression correlating with different life history stages or cell types. Notably, two ESTs encoding HD proteins showed eightfold higher expression in the P. purpurea sporophyte (conchocelis) than in the gametophyte (blade), whereas two MADS domain-containing paralogs showed significantly different patterns of expression in the conchocelis and blade respectively. These developmental gene families do not appear to have undergone the kinds of dramatic expansions in copy number found in multicellular land plants and animals, which are important for regulating developmental processes in those groups. Analyses of small RNAs did not validate the presence of miRNAs, but homologs of Argonaute were present. In general, it appears that red algae began with a similar molecular toolkit for directing development as did other multicellular eukaryotes, but probably evolved altered roles for many key proteins, as well as novel mechanisms yet to be discovered.
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: John W
authors.initials: JW
authors.lastname: Stiller
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Justin
authors.initials: J
authors.lastname: Perry
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Linda A
authors.initials: LA
authors.lastname: Rymarquis
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Monica
authors.initials: M
authors.lastname: Accerbi
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Pamela J
authors.initials: PJ
authors.lastname: Green
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Simon
authors.initials: S
authors.lastname: Prochnik
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Erika
authors.initials: E
authors.lastname: Lindquist
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Cheong Xin
authors.initials: CX
authors.lastname: Chan
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Charles
authors.initials: C
authors.lastname: Yarish
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Senjie
authors.initials: S
authors.lastname: Lin
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Yunyun
authors.initials: Y
authors.lastname: Zhuang
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Nicolas A
authors.initials: NA
authors.lastname: Blouin
authors.affiliation: Department of Biology, East Carolina University, Greenville, NC 27848, USADelaware Biotechnology Institute, Delaware Technology Park, Newark DE 19711, USADOE Joint Genomics Institute, Walnut Creek, CA 94598, USADepartment of Ecology, Evolution and Natural Resources, Rutgers University, New Brunswick, NJ 08901, USADepartment of Ecology and Evolutionary Biology, University of Connecticut, Stamford, CT, 06901, USADepartment of Marine Sciences, University of Connecticut, Groton, CT 06340, USASchool of Marine Science, University of Maine, Orono, ME 04469 USA.
authors.forename: Susan H
authors.initials: SH
authors.lastname: Brawley
keywords: EST; MADS; Porphyra; Rhodophyta; SWI/SNF; development; evolution; homeodomain; miRNA; small RNA