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Provided for Non-Commercial Research and Educational Use. Not for Reproduction, Distribution Or Commercial Use Provided for non-commercial research and educational use. Not for reproduction, distribution or commercial use. This article was originally published in the Encyclopedia of Evolutionary Biology published by Elsevier, and the attached copy is provided by Elsevier for the author's benefit and for the benefit of the author's institution, for non- commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues who you know, and providing a copy to your institution's administrator. All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution's website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier's permissions site at: http://www.elsevier.com/locate/permissionusematerial Gontier, N. (2016) Symbiogenesis, History of. In: Kliman, R.M. (ed.), Encyclopedia of Evolutionary Biology. vol. 4, pp. 261–271. Oxford: Academic Press. © 2016 Elsevier Inc. All rights reserved. Author's personal copy Symbiogenesis, History of N Gontier, University of Lisbon, Lisbon, Portugal r 2016 Elsevier Inc. All rights reserved. Glossary Horizontal transmission Any type of exchange between Archaea Oldest Domain of Life, previously designated as distinct individuals that happens during their lifetime and Archaebacteria. outside of the germ line. Bacteria Second Domain of Life, previously designated as Kingdoms of Life/ five-kingdom classification According Eubacteria. to Whittaker and Margulis (1978), life can be classified into Domains of life/ three-domain classification Based five kingdoms: Prokaryotic Monera (containing the upon results obtained from comparative molecular Archaebacteria and Eubacteria), Eukaryotic Protoctista phylogenetics, Woese et al. (1990) undid the prokaryote/ (Protists), Fungi, Plants, and Animals. eukaryote and five-kingdom classification and instead Lateral gene transfer Horizontal gene exchange between distinguished three major domains of life: Archaea, Bacteria, distinct organisms or distinct genomes within the same and Eukaryota. organism. Eukaryota/Eukaryotes Third domain of life, containing Metamorphosis Pre-evolutionary idea that living the Protist, Fungi, Plant and Animal Kingdoms. Eukaryotic organisms can transform, or change in form. cells have a membrane-bounded nucleus that contains the Transformation and metamorphosis are precursors to genome, and the cytoplasm often contains various cellular transmutation and evolutionary theory. bodies called organelles. Microbiome The complete community of microorganisms Flagellum/Flagella Whip-like extensions of prokaryotic that inhabit or live on the surface of an organism. The term cells that enable motility. Contrary to eukaryotic is sometimes used to specifically refer to the genomes of the undulipodia, flagella are made up of flagelin proteins. microbiota. Hereditary symbiosis Symbiotic association that becomes Microtubules Tubulin protein structures. permanent and irreversible, foundational for Prokaryotes All Archaea and Bacteria, typified by a free- symbiogenesis. floating instead of nucleated genome. Defining Symbiogenesis wherefrom mitochondria and chloroplasts evolved still exist as free-living bacteria today. Symbiosis occurs when distinct organisms live in close asso- That mitochondria and chloroplasts evolved by symbiogen- ciation with one another, while symbiogenesis is both a phe- esis has for the most part of history been suggested based upon nomenon and an evolutionary mechanism (Merezhkowsky, morphological comparisons between the organelles and bacteria, 1905, 1910; Famintsyn, 1907; Kozo-Polyansky, 1924/2010) as well as the fact that these organelles contain their own DNA that results from permanent and hereditary symbiosis (von and have a double membrane, suggestive of bacterial engulfment Faber, 1912; Buchner, 1921, 1939; Wallin, 1927; Lederberg, (Ris and Plaut, 1962; Nass and Nass, 1963). Genetic com- 1952; Sagan, 1967). Margulis and Dolan (2000, p. 157) define parisons between the organellar DNA and the DNA of bacterial symbiogenesis as the “origin of a new organ, metabolic lineages has now confirmed their bacterial origin. Chloroplasts pathway, behavior, tissue, or other feature as a result of long- evolved from cyanobacteria, and mitochondria evolved from term hereditary symbiosis and includes the process by which proteobacteria (Bonen and Doolittle, 1975; Bonen and Doolittle, organelles of the eukaryotic cell evolved from ancient bacterial 1976; Bonen et al., 1977; Gray and Doolittle, 1982). Molecular symbionts.” gene-sequencing techniques have furthermore shown instances Symbiogenesis today is well recognized to underlie the of lateral gene transfer between the organelles and the nucleus, in origin of mitochondria and chloroplasts. Both are cellular both directions. After symbiogenetic acquisition, mitochondrial organelles (organ-like bodies), found exclusively in eukary- and chloroplast DNA underwent considerable gene loss otic life forms (prokaryotes lack them, Figure 1), where they (Archibald, 2014; Martin and Herrmann, 1998). reside inside the cytoplasm of the cell (Figure 2). Mito- Eukaryotic cells often have many more organelles, and their chondria are present in aerobe protist, plant, and animal evolutionary origin remains uncertain. A bacterial and sym- cells, while algal and plant cells also contain chloroplasts. biogenetic origin for the lysosomes has been suggested by its These organelles once used to be free-living bacteria that, discoverer, the Belgian cytologist De Duve et al. (1974). around 2 billion years ago, entered some of the first eu- According to Margulis, the nucleated cell also evolved by karyotic life forms through phagocytosis (eating or engulf- means of symbiogenesis. As such, it was symbiogenesis that ment). The organisms engaged in symbiosis, this became enabled both the evolutionary transition from prokaryotes to permanent and hereditary, and the once free-living organisms eukaryotes, and the subsequent evolution of the four eukary- evolved into the organelles (Figure 3). The bacterial lineages otic kingdoms. Encyclopedia of Evolutionary Biology, Volume 4 doi:10.1016/B978-0-12-800049-6.00016-0 261 Author's personal copy 262 Symbiogenesis, History of Translation of RNA into proteins via ribosomes Singular RNA Transcription of DNA into RNA via RNA polymerase Amino acids Circular Ribosomes double Plasma membrane stranded DNA Micro- Cell wall compartments Capsule (optional) Flagellum (optional) Figure 1 Schematic of a prokaryote. Prokaryotes neither have a membrane-bounded nucleus nor cellular organelles though they often contain micro-compartments that package enzymes and proteins. From Hereditary Symbiosis to Symbiogenesis: The All assumed and still assume today, that one plasma underlies all Origin of Mitochondria and Chloroplasts organisms, in other words, that out of non-being, life came forth from one root, from where one tree of organisms developed, first as a common trunk of protists, and then the tree split into two main axes Symbiosis research was often conducted in the margins of – the plant axis and the animal axis. Until now, there was the general Darwinian evolutionary biology. Boveri (1904), for example, conviction, that the tree of life was a single one. The task set forth in although a founder of chromosome theory, merely stated that this work, is to demonstrate that there are two trees of life, and that each tree originated on its own and independently from the other Mendelian factors are transmitted via chromosomes. He also ’ ‘ ’ one, and this probably happened in different periods of earth s conjectured that the protoplasm (cytoplasm) and chromo- history. These trees partly developed on their own and independ- somes of the cell originated through symbiosis, an idea that ently from one another and partly stringed together and closely grew was already introduced in 1893 by Shôsaburô Watasé (Sapp, and developed together. Both trees are responsible for the diversity 1994; Carrapiço, 2010, 2015). of organic beings. The idea of a unity of organic nature has to be abandoned in favor of the idea of nature’s duality. (Merezhkowsky, Earlier in time, Schimper (1883, p. 112) suggested a single 1910, my translation) origin for “Chlorophyllkörner” (chlorophyll grains) and “Farb- körper” (pigment corpuscles) present in the plastids of plant and algal cells (“plastiden,” subdivided into colorless “leukoplastiden,” green “chloroplastiden” and yellow, orange and red “chromoplasti- Life evolved from two distinct organismal types, each den”). He noted that they never arise de novo but develop from consisting of different ‘protoplasms’ (“plasma-arten”), ’’Myko- pre-existing ‘protoplasmic’ structures, and he thought they could plasma’’ (Mycoplasma) and ’’Amöboides Plasma’’ (amoeba-like divide and ‘metamorphose’ into one another. Because their plasma). Mykoplasma is anaerobe, autotrophic, rich in phos- protein structures (“Proteïnkrystalle”) resemble those of ‘living phor and nucleic acid (’’nuclein’’), and experiments demon- plasma’ (bacteria), Schimper (1885, p. 202) later conjectured strated a heat tolerance of up to 90 1C and a high resistance to that chloroplasts might have resulted from a symbiosis between a poisons. Amöboplasma is aerobe and heterotrophic, low in colorless and chlorophyll-containing organism. phosphor and nucleic acids, it can only bear temperatures
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