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The Third International Colloquium on Endocytobiology June 10-12, 1986 The World Trade Center New York City • Meeting presided over by John J. Lee and Jerome F. Fredrick, both on the faculty at City University of New York • Presentations related to the interrelationships between living things: all types of symbioses Lee: 1933- Fredrick: 1926-1995 Some Background on Endosymbiosis Endosymbiotic origin of chloroplasts and mitochondria proposed much earlier. Endosymbiotic Origin of Eukaryotes Schimper (1883) and Mereschcowsky (1905) proposed that chloroplasts are cyanobacteria living inside plant cells Andreas Franz Wilhelm Schimper (1856-1901, Germany) Konstantin Sergeevich Mereschcowsky (1855-1921, Russia) Mereschcowsky, a lichenologist, rejected the importance of natural selection and explained evolutionary novelties by acquisition of bacteria through symbiosis. Mereschcowsky’s tree of life, 1910 Ivan Emanuel Wallin • Rejected cytoplasmic origin of mitochondria • Considered them to be microbes living in the cytoplasm • Claimed to have cultured isolated mitochondria as proof of their microbial nature 1883-1969, USA Lynn Sagan (1967) • On the Origin of Mitosing Cells. J. Theoret. Biol. 14: 225-274. • Proposed the 9+2 basal body together with mitochondria and chloroplasts was an endosymbiotic microbe • Basal body from a spirochaete, a microbe that has multiple microbial flagella (microtubules of flagellin) between the two membranes. • Microtubular array extended inward makes spindle and outward becomes a flagellum • Basal bodies also function as centrioles • Endosymbiosis established as a working theory by 1986. • Examples of symbiosis and endosymbiosis on program • Models of organisms that meet criteria for the endosymbiotic process (ex: Thermoplasma, an archean without a wall as a model for the nuclear host) Mixotricha paradoxa • Trichomonad that inhabits gut of Australian termite. • Has two species of surface bacterial symbionts: spirochaetes provide locomotion and serve as de facto flagella • Internal mitochondria-like symbionts • Together with the nucleus and the genome of the hydrogenosome (Hproducing ‘degenerate’ mitochondrion), they function together as 5 genomes. Frank Round (1927-2010, Britain) Proposal by Frank Round (1980). Endosymbiosis would make evolutionary history of eukaryotes impossible to determine Lynn (Sagan) Margulis (1938-2011, USA) • Scientific iconoclast • Founder of modern endosymbiotic theory • Co-Founder of Gaia Hypothesis • Chief apologist for the 5kingdom system • After 2008 – AIDS due to syphilis – 9/11 conspiracy theorist Receiving National Medal of Science 2000 F. J. R. Max Taylor Serial endosymbiotic theory (SET (19741990)) organelles are the result of successive engulfments… Max F.J.R. Taylor (1939, South Africa and Canada); eukaryote created following endosymbiosis with mitochondrial bacterium. Further developed Margulis Endosymbiosis Taylor (1976) Ran counter to Margulis (1967) in that organisms without flagella were most primitive –the prevailing scientific view Thomas Cavalier-Smith • Proposed eukaryotes evolved gradually from bacteria which lost the wall and relied on cytoskeleton for cellular integrity • The internal microtubular array gave rise to flagella, mitotic spindles, etc. 1942- Britain; Professor emeritus, University of Oxford Cavalier-Smith (2010) Figure 3 from: Philos. Trans. R. Soc. Lond. B Biol. Sci. 365(1537): 111–132. The Dispersed Archaeal Eukaryome and the Complex Archaeal Ancestor of Eukaryotes Eugene V. Koonin and Natalya Yutin (2014) • Host derived from Crenarchaea • Sophisticated cytoskeleton • Phagocytic capabilities • No wall • Mesophylic • Co-existed with other bacteria • Capture many genes through HGT (LTG) Archaea or Archaeon? Endosymbiotic theory for organelle origins Verena Zimorski, Chuan Ku, William F Martin and Sven B Gould (2014) Endosymbiotic theory for organelle origins Verena Zimorski, Chuan Ku, William F Martin and Sven B Gould (2014) Theories of the Origin of the Nucleus Martin et al. (2015) The Inside-Out Model for the Origin of the Nucleus (Baum and Baum 2014) Origin of Cell Division (Baum and Baum 2014) Life Ascending Nick Lane (2009) Complex life not inevitable – Accident of one getting inside another – Accident of a selfish genome gathering genes by HGT from environment and symbiont then using barriers in the cell to slow down mRNA to splice introns – These accidents allowed the new cells to be larger in size and apply a concomitantly larger set of physiological pathways Reader in Evolutionary Biochemistry in the Department of Genetics, Evolution and Environment at University College London Power from Chemiosmosis drives complexity (Lane and Martin 2010)