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== Evolutionary origins == The protein content of peroxisomes varies across species or organism, but the presence of proteins common to many species has been used to suggest an [[Endosymbiotic theory|endosymbiotic]] origin; that is, peroxisomes evolved from bacteria that invaded larger cells as parasites, and very gradually evolved a symbiotic relationship.<ref name="pmid3916321">{{cite journal | vauthors = Lazarow PB, Fujiki Y | title = Biogenesis of peroxisomes | journal = Annual Review of Cell Biology | volume = 1 | pages = 489–530 | year = 1985 | pmid = 3916321 | doi = 10.1146/annurev.cb.01.110185.002421 }}</ref> However, this view has been challenged by recent discoveries.<ref name="pmid17506702">{{cite journal | vauthors = Fagarasanu A, Fagarasanu M, Rachubinski RA | title = Maintaining peroxisome populations: a story of division and inheritance | journal = Annual Review of Cell and Developmental Biology | volume = 23 | pages = 321–44 | year = 2007 | pmid = 17506702 | doi = 10.1146/annurev.cellbio.23.090506.123456 }}</ref> For example, peroxisome-less mutants can restore peroxisomes upon introduction of the wild-type gene. Two independent evolutionary analyses of the peroxisomal [[proteome]] found homologies between the peroxisomal import machinery and the [[Endoplasmic Reticulum Associated Protein Degradation|ERAD]] pathway in the [[endoplasmic reticulum]],<ref name="pmid16452116">{{cite journal | vauthors = Schlüter A, Fourcade S, Ripp R, Mandel JL, Poch O, Pujol A | title = The evolutionary origin of peroxisomes: an ER-peroxisome connection | journal = Molecular Biology and Evolution | volume = 23 | issue = 4 | pages = 838–45 | date = April 2006 | pmid = 16452116 | doi = 10.1093/molbev/msj103 | doi-access = free }}</ref><ref name="pmid16556314">{{cite journal | vauthors = Gabaldón T, Snel B, van Zimmeren F, Hemrika W, Tabak H, Huynen MA | title = Origin and evolution of the peroxisomal proteome | journal = Biology Direct | volume = 1 | pages = 8 | date = March 2006 | pmid = 16556314 | pmc = 1472686 | doi = 10.1186/1745-6150-1-8 | doi-access = free }}</ref> along with a number of metabolic enzymes that were likely recruited from the [[mitochondria]].<ref name="pmid16556314" /> The peroxisome may have had an [[Actinomycetota]] origin;<ref name="pmid19818387">{{cite journal | vauthors = Duhita N, Le HA, Satoshi S, Kazuo H, Daisuke M, Takao S | title = The origin of peroxisomes: The possibility of an actinobacterial symbiosis | journal = Gene | volume = 450 | issue = 1–2 | pages = 18–24 | date = January 2010 | pmid = 19818387 | doi = 10.1016/j.gene.2009.09.014 }}</ref> however, this is controversial.<ref name="pmid20600706">{{cite journal | vauthors = Gabaldón T, Capella-Gutiérrez S | title = Lack of phylogenetic support for a supposed actinobacterial origin of peroxisomes | journal = Gene | volume = 465 | issue = 1–2 | pages = 61–5 | date = October 2010 | pmid = 20600706 | doi = 10.1016/j.gene.2010.06.004 }}</ref>
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