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Algal Evolution Johanna Weston Keeling, 2004
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3.5 billion cyanobacteria 2.2 to 1.5 billion mitochondial origin 1.5 – 1.2 Plastid “Little Green Slaves” (Dyall et al., 2004)
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ENDOSYMBIOSIS – Primary – Secondary – Tertiary – Serial Secondary (Keeling, 2004)
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How do endosymbionts become organelles? Organelle – discrete subcellular structure of specialized function usually bound by two or more membranes Metabolite antiporters and Biochemical pathways Genetic integration and reduction Protein import apparatus
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Phosphate Translocator Family Antiport dependent on counter-exchange Algae profit from cyanobacteria carbon fixation ER/Golgi metabolite translocator protein in host recruited to the plastid envelope (Bhattacharya et al., 2007)
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Transfer and Reduction Cyanobacteria – 2000 to 4000 kb Plastids – < 200 kb – < 200 genes “Muller’s ratchet” Oxygen free radials (Bhattacharya et al., 2007)
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TIC - TOC Defining feature of organelle genesis Sophisticated outcome of the requirement for regulated protein import Evolutionary diverse origins – Cyanobacteria – Co-option of host genes – HGT from bacteria Transit peptideN C (Gould et al., 2008)
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TIC-TOC Independent Existing endomembrane system of host cell Paulinella chromatophora α – carbonic anhydrase **2 o Endosymbiosis combination of both (Gould et al., 2008)(Bhattacharya et al., 2007)
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Animal-Algal Endosymbiosis Elysia chlorotica and chloroplasts of Vaucheria litorea 9 months with only light and CO 2 Not inherited (Mujer et al., 1996) (Green et al., 2000)
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References Bhattacharya, Debashish, et al. "How do endosymbionts become organelles? Understanding early events in plastid evolution." BioEssay (2007): 1239-1246. Gould, Sven, Ross Waller and Geoffrey Macfadden. "Plastid evolution." Annual Review of Plant Biology (2008): 491-517. Graham, Linda E., James M. Graham and Lee W. Wilcox. Algae. 2. San Francisco: Benjamin Cummings, 2009. Green, Brian, et al. "Mullusc-algal chloroplast endosymbiosis. Photosynthesis, thylakoid protein maintenance, and chloroplast gene expression continue for many months in the absence of the algal nucleus." Plant Physiology (2000): 331-342. Keeling, Patrick. "Diversity and evolutionary history of plastids and their hosts." Americal Journal of Botany (2004): 1418-1493. Mujer, Cesar, et al. "Chloroplast genes are expressed during intracellular symbiotic association of Vaucheria litorea plastids with the sea slug Elysia chlorotica." Cell Biology (1996): 12333-12338. Weber, Andreas, Marc Linka and Debashish Bhattacharya. "Single, ancient orgin of a plastid metabolite translocator family in Plantae from an endomembrane-derived ancestor." Eukaryotic Cell (2006): 609-612.
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