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Nucleomorph
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==Organisms with known nucleomorphs== As of 2007, only two [[Monophyly|monophyletic]] groups of organisms are known to contain plastids with a vestigial nucleus or nucleomorph: the [[cryptomonads]]<ref name=Lane2007>{{Cite journal | last1 = Lane | first1 = C. E. | last2 = Van Den Heuvel | first2 = K. | last3 = Kozera | first3 = C. | last4 = Curtis | first4 = B. A. | last5 = Parsons | first5 = B. J. | last6 = Bowman | first6 = S. | last7 = Archibald | first7 = J. M. | doi = 10.1073/pnas.0707419104 | title = Nucleomorph genome of Hemiselmis andersenii reveals complete intron loss and compaction as a driver of protein structure and function | journal = Proceedings of the National Academy of Sciences | volume = 104 | issue = 50 | pages = 19908β19913 | year = 2007 | pmid = 18077423 | pmc =2148396 | bibcode = 2007PNAS..10419908L| doi-access = free }}</ref> of the supergroup [[Cryptista]] and the [[chlorarachniophyte]]s<ref name=Gilson2006>{{Cite journal | last1 = Gilson | first1 = P. R. | last2 = Su | first2 = V. | last3 = Slamovits | first3 = C. H. | last4 = Reith | first4 = M. E. | last5 = Keeling | first5 = P. J. | last6 = McFadden | first6 = G. I. | title = Complete nucleotide sequence of the chlorarachniophyte nucleomorph: Nature's smallest nucleus | doi = 10.1073/pnas.0600707103 | journal = Proceedings of the National Academy of Sciences | volume = 103 | issue = 25 | pages = 9566β9571 | year = 2006 | pmid = 16760254 | pmc =1480447 | bibcode = 2006PNAS..103.9566G| doi-access = free }}</ref> of the supergroup [[Rhizaria]], both of which have examples of sequenced nucleomorph genomes.<ref name=Lane2007 /><ref name=Gilson2006 /> Studies of the genomic organization and of the [[molecular phylogeny]] have shown that the nucleomorph of the [[cryptomonads]] used to be the [[Cell nucleus|nucleus]] of a [[red algae|red alga]], whereas the nucleomorph of the chlorarchniophytes was the nucleus of a [[green algae|green alga]]. In both groups of organisms the plastids originate from engulfed [[photoautotroph]]ic [[eukaryote]]s. Of the two known plastids that contain nucleomorphs, both have four membranes, the nucleomorph residing in the [[periplastidial compartment]], evidence of being engulfed by a eukaryote through [[phagocytosis]].<ref name=archibald1/> In 2020, genetic work identified the plastid in ''[[Lepidodinium]]'' and two previously undescribed dinoflagellates ("MGD" and "TGD") as being most closely related to the green alga ''[[Pedinomonas]]''. The observation of a nucleomorph in ''Lepidodinium'' is controversial, but MGD and TGD are proven to have DNA-containing nucleomorphs.<ref name=Pedinomonas>{{cite journal |last1=Sarai |first1=C |last2=Tanifuji |first2=G |last3=Nakayama |first3=T |last4=Kamikawa |first4=R |last5=Takahashi |first5=K |last6=Yazaki |first6=E |last7=Matsuo |first7=E |last8=Miyashita |first8=H |last9=Ishida |first9=KI |last10=Iwataki |first10=M |last11=Inagaki |first11=Y |title=Dinoflagellates with relic endosymbiont nuclei as models for elucidating organellogenesis. |journal=Proceedings of the National Academy of Sciences of the United States of America |date=10 March 2020 |volume=117 |issue=10 |pages=5364β5375 |doi=10.1073/pnas.1911884117 |pmid=32094181 |doi-access=free|pmc=7071878 |bibcode=2020PNAS..117.5364S }}</ref> The transcriptomes of the nucleomorphs have been sequenced.<ref>[https://www.tandfonline.com/doi/pdf/10.1080/19420889.2020.1776568 Putative genome features of relic green alga-derived nuclei in dinoflagellates and future perspectives as model organisms]</ref> One slight issue in understanding the sequence of evolution is that although the phylogenetic tree built from Lepidodinium-MGD-TGD's plastid is monophyletic, the tree built from their host-nucleus DNA is not, implying that they might have acquired very similar algae independently.<ref name=Pedinomonas/> <gallery> File:2023 Cryptomonad.svg|Representation of a Cryptomonad File:2023 Chlorarachniophyte.svg|Representation of a Chlorarachniophyte </gallery>
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