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==Role in evolution== [[File:Common jassid nymphs and ants02.jpg|thumb|upright|[[Leafhopper]]s protected by [[meat ant]]s]] {{Further|Co-evolution}} Symbiosis is increasingly recognized as an important selective force behind evolution;<ref name="Moran-2006" /><ref>{{Harvnb|Wernegreen|2004}}</ref> many species have a long history of interdependent [[co-evolution]].<ref>{{Harvnb|Paracer|Ahmadjian|2000|pp=3–4}}</ref> Although symbiosis was once discounted as an anecdotal evolutionary phenomenon, evidence is now overwhelming that obligate or facultative associations among microorganisms and between microorganisms and multicellular hosts had crucial consequences in many landmark events in evolution and in the generation of phenotypic diversity and complex phenotypes able to colonise new environments.<ref name="Mutalipassi-2021">{{cite journal |last1=Mutalipassi |first1=Mirko |last2=Riccio |first2=Gennaro |last3=Mazzella |first3=Valerio |last4=Galasso |first4=Christian |last5=Somma |first5=Emanuele |last6=Chiarore |first6=Antonia |last7=de Pascale |first7=Donatella |last8=Zupo |first8=Valerio |display-authors=3 |title=Symbioses of Cyanobacteria in Marine Environments: Ecological Insights and Biotechnological Perspectives |journal=Marine Drugs |volume=19 |issue=4 |page=227 |date=April 2021 |pmid=33923826 |pmc=8074062 |doi=10.3390/md19040227 |publisher=MDPI AG |doi-access=free }} [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].</ref> [[Mutualism (biology)|Mutualistic]] symbiosis can sometimes evolve from [[parasitism]] or [[commensalism]], Fungi's relationship to plants in the form of [[mycelium]] evolved from parasitism and [[commensalism]]. Under certain conditions species of fungi previously in a state of mutualism can turn parasitic on weak or dying plants.<ref>{{cite journal |last1=Purin |first1=Sonia |last2=Rillig |first2=Matthias C. |title=Parasitism of arbuscular mycorrhizal fungi: reviewing the evidence |journal=FEMS Microbiology Letters |date=February 2008 |volume=279 |issue=1 |pages=8–14 |doi=10.1111/j.1574-6968.2007.01007.x |pmid=18070075 }}</ref> Likewise the symbiotic relationship of [[clown fish]] and [[sea anemone]]s emerged from a commensalist relationship.<ref>{{Cite web|url=https://yaledailynews.com/blog/2018/02/06/yale-researchers-show-that-mutualism-can-come-from-parasitism/#:~:text=The+viruses+and+bacteria+originally,benefited+from+one+another's+presence|title=Yale researchers show that mutualism can come from parasitism|first=Jessica|last=Pevner|date=February 6, 2018|website=Yale Daily News}}</ref><ref>{{cite journal |last1=Titus |first1=Benjamin M. |last2=Benedict |first2=Charlotte |last3=Laroche |first3=Robert |last4=Gusmão |first4=Luciana C. |last5=Van Deusen |first5=Vanessa |last6=Chiodo |first6=Tommaso |last7=Meyer |first7=Christopher P. |last8=Berumen |first8=Michael L. |last9=Bartholomew |first9=Aaron |last10=Yanagi |first10=Kensuke |last11=Reimer |first11=James D. |last12=Fujii |first12=Takuma |last13=Daly |first13=Marymegan |last14=Rodríguez |first14=Estefanía |title=Phylogenetic relationships among the clownfish-hosting sea anemones |journal=Molecular Phylogenetics and Evolution |date=October 2019 |volume=139 |pages=106526 |doi=10.1016/j.ympev.2019.106526 |pmid=31158486 |bibcode=2019MolPE.13906526T }}</ref><ref>{{cite journal |last1=Litsios |first1=Glenn |last2=Sims |first2=Carrie A |last3=Wüest |first3=Rafael O |last4=Pearman |first4=Peter B |last5=Zimmermann |first5=Niklaus E |last6=Salamin |first6=Nicolas |title=Mutualism with sea anemones triggered the adaptive radiation of clownfishes |journal=BMC Evolutionary Biology |date=December 2012 |volume=12 |issue=1 |page=212 |doi=10.1186/1471-2148-12-212 |doi-access=free |pmid=23122007 |pmc=3532366 |bibcode=2012BMCEE..12..212L }}</ref> === Hologenome development and evolution === Evolution originated from changes in development where variations within species are selected for or against because of the symbionts involved.<ref name="Li-2015">{{cite journal |last1=Li |first1=Ci-Xiu |last2=Shi |first2=Mang |last3=Tian |first3=Jun-Hua |last4=Lin |first4=Xian-Dan |last5=Kang |first5=Yan-Jun |last6=Chen |first6=Liang-Jun |last7=Qin |first7=Xin-Cheng |last8=Xu |first8=Jianguo |last9=Holmes |first9=Edward C |last10=Zhang |first10=Yong-Zhen |display-authors=3 |title=Unprecedented genomic diversity of RNA viruses in arthropods reveals the ancestry of negative-sense RNA viruses |journal=eLife |volume=4 |issue=|pages=|date=January 2015 |pmid=25633976 |pmc=4384744 |doi=10.7554/eLife.05378 |url=|doi-access=free }}</ref> The hologenome theory relates to the holobiont and symbionts genome together as a whole.<ref>{{cite journal |last1=Rosenberg |first1=E. |last2=Zilber-Rosenberg |first2=I. |title=Symbiosis and development: the hologenome concept |journal=Birth Defects Research. Part C, Embryo Today |volume=93 |issue=1 |pages=56–66 |date=March 2011 |pmid=21425442 |doi=10.1002/bdrc.20196 }}</ref> Microbes live everywhere in and on every multicellular organism.<ref>{{cite journal |last=Morris |first=J.J. |title=What is the hologenome concept of evolution? |journal=F1000Research |volume=7 |pages=1664 |date=2018-10-19 |pmid=30410727 |pmc=6198262 |doi=10.12688/f1000research.14385.1 |doi-access=free }}</ref> Many organisms rely on their symbionts in order to develop properly, this is known as co-development. In cases of co-development the symbionts send signals to their host which determine developmental processes. Co-development is commonly seen in both arthropods and vertebrates.<ref name="Li-2015"/> === Symbiogenesis === {{main|Symbiogenesis}} One hypothesis for the origin of the nucleus in [[eukaryotes]] (plants, animals, fungi, and [[protists]]) is that it developed from a [[symbiogenesis]] between bacteria and archaea.<ref name="Moran-2006"/><ref>{{Harvnb|Brinkman|Blanchard|Cherkasov|Av-Gay|2002}}</ref><ref>{{Harvnb|Golding|Gupta|1995}}</ref> It is hypothesized that the symbiosis originated when ancient archaea, similar to modern methanogenic archaea, invaded and lived within bacteria similar to modern myxobacteria, eventually forming the early nucleus. This theory is analogous to the accepted theory for the origin of eukaryotic mitochondria and chloroplasts, which are thought to have developed from a similar endosymbiotic relationship between proto-eukaryotes and aerobic bacteria.<ref>{{cite book |last=Margulis |first=Lynn |author-link=Lynn Margulis |year=1981 |title=Symbiosis in Cell Evolution |pages=[https://archive.org/details/symbiosisincelle00marg/page/206 206–227] |publisher=W. H. Freeman and Company |location=San Francisco |isbn=978-0-7167-1256-5 |url=https://archive.org/details/symbiosisincelle00marg/page/206 }}</ref> Evidence for this includes the fact that [[mitochondria]] and [[chloroplast]]s divide independently of the cell, and that these organelles have their own genome.<ref>"Symbiosis". ''Bloomsbury Guide to Human Thought''. London: Bloomsbury Publishing, 1993. Credo Reference. Web. 17 September 2012.</ref> The biologist [[Lynn Margulis]], famous for her work on [[endosymbiosis]], contended that symbiosis is a major driving force behind [[evolution]]. She considered [[Charles Darwin|Darwin]]'s notion of evolution, driven by competition, to be incomplete and claimed that evolution is strongly based on [[Co-operation (evolution)|co-operation]], [[Biological interaction|interaction]], and [[mutual dependence]] among organisms. According to Margulis and her son [[Dorion Sagan]], "[[Life]] did not take over the [[Earth|globe]] by [[combat]], but by [[social network|networking]]."<ref>{{harvnb|Sagan|Margulis|1986}}</ref>
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