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@ARTICLE{ReineckeLevi:868043,
      author       = {Reinecke-Levi, Diana and Zarka, Aliza and Leu, Stefan and
                      Boussiba, Sammy},
      title        = {{C}loning, molecular characterization, and phylogeny of two
                      evolutionary distinct glutamine synthetase isoforms in the
                      green microalga {H}aematococcus pluvialis ({C}hlorophyceae)},
      journal      = {Journal of phycology},
      volume       = {52},
      number       = {6},
      issn         = {0022-3646},
      address      = {Oxford [u.a.]},
      publisher    = {Wiley-Blackwell},
      reportid     = {FZJ-2019-06636},
      pages        = {961 - 972},
      year         = {2016},
      abstract     = {Haematococcus pluvialis (Chlorophyta) is a widely used
                      microalga of great economic potential, yet its molecular
                      genetics and evolution are largely unknown. We present new
                      detailed molecular and phylogenetic analysis of two
                      glutamine synthetase (GS) enzymes and genes (gln) under the
                      Astaxanthin‐inducing conditions of light‐ and
                      nitrogen‐stress. Structure analysis identified key
                      residues and confirmed two decameric GS2 holoenzymes, a
                      cytoplasmic enzyme, termed GS2c, and a plastidic form,
                      termed GS2p, due to chloroplast‐transit peptides at its
                      N‐terminus. Gene expression analysis showed dissociation
                      of mRNA, protein, and enzyme activity levels for both GS2
                      under different growth conditions, indicating the strong
                      post‐transcriptional regulation. Data‐mining identified
                      novel and specified published gln genes from Prasinophyceae,
                      Chlorophyta, Trebouxiophyceae, Charophyceae, Bryophyta,
                      Lycopodiophyta, Spermatophyta, and Rhodophyta. Phylogenetic
                      analysis found homologues to the cytosolic GS2c of H.
                      pluvialis in all other photo‐ and non‐photosynthetic
                      Eukaryota. The chloroplastic GS2p was restricted to
                      Chlorophyta, Bryophyta, some Proteobacteria and Fungii; no
                      homologues were identified in Spermatophyta or other
                      Eukaryota. This indicates two independent prokaryotic donors
                      for these two gln genes in H. pluvialis. Combined
                      phylogenetic analysis of GS, chl‐b synthase, elongation
                      factor, and light harvesting complex homologues project a
                      newly refined model of Viridiplantae evolution. Herein, a
                      GS1 evolved into the cytosolic GS2c and was passed on to all
                      Eukaryota. Later, the chloroplastic GS2p entered the
                      Archaeplastida lineage via a horizontal gene transfer at the
                      divergence of Chlorophyta and Rhodophyta lineages. GS2p
                      persisted in Chlorophyta and Bryophyta, but was lost during
                      Spermatophyta evolution. These data suggest the revision of
                      GS classification and nomenclature, and extend our
                      understanding of the photosynthetic Eukaryota evolution.},
      cin          = {IBG-2},
      ddc          = {580},
      cid          = {I:(DE-Juel1)IBG-2-20101118},
      pnm          = {582 - Plant Science (POF3-582)},
      pid          = {G:(DE-HGF)POF3-582},
      typ          = {PUB:(DE-HGF)16},
      pubmed       = {pmid:27402429},
      UT           = {WOS:000390344600007},
      doi          = {10.1111/jpy.12444},
      url          = {https://juser.fz-juelich.de/record/868043},
}