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Cell nucleus
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==Nuclei per cell== Most [[eukaryotic]] cell types usually have a single nucleus, but some have no nuclei, while others have several. This can result from normal development, as in the maturation of mammalian [[red blood cell]]s, or from faulty cell division.<ref>{{cite book |last1=Ressel |first1=Lorenzo | name-list-style = vanc | chapter = Nuclear Morphologies |title=Normal cell morphology in canine and feline cytology: an identification guide |date=2017 |publisher=John Wiley & Sons |location=Hoboken, NJ |isbn=978-1-119-27891-7 |page=6 | chapter-url=https://books.google.com/books?id=AkwrDwAAQBAJ&pg=PA6}}</ref> ===Anucleated cells=== [[Image:redbloodcells.jpg|thumb|230px|right|Human red blood cells, like those of other mammals, lack nuclei. This occurs as a normal part of the cells' development.]] An anucleated cell contains no nucleus and is, therefore, incapable of dividing to produce daughter cells. The best-known anucleated cell is the mammalian red blood cell, or [[erythrocyte]], which also lacks other organelles such as mitochondria, and serves primarily as a transport vessel to ferry [[oxygen]] from the [[lungs]] to the body's tissues. Erythrocytes mature through [[erythropoiesis]] in the [[bone marrow]], where they lose their nuclei, organelles, and ribosomes. The nucleus is expelled during the process of differentiation from an [[erythroblast]] to a [[reticulocyte]], which is the immediate precursor of the mature erythrocyte.<ref name="Skutelsky">{{cite journal | vauthors = Skutelsky E, Danon D | title = Comparative study of nuclear expulsion from the late erythroblast and cytokinesis | journal = Experimental Cell Research | volume = 60 | issue = 3 | pages = 427β36 | date = June 1970 | pmid = 5422968 | doi = 10.1016/0014-4827(70)90536-7 | department = Primary }}</ref> The presence of [[mutagen]]s may induce the release of some immature "micronucleated" erythrocytes into the bloodstream.<ref name="Torous">{{cite journal | vauthors = Torous DK, Dertinger SD, Hall NE, Tometsko CR | title = Enumeration of micronucleated reticulocytes in rat peripheral blood: a flow cytometric study | journal = Mutation Research | volume = 465 | issue = 1β2 | pages = 91β9 | date = February 2000 | pmid = 10708974 | doi = 10.1016/S1383-5718(99)00216-8 | bibcode = 2000MRGTE.465...91T | department = Primary }}</ref><ref name="Hutter">{{cite journal | vauthors = Hutter KJ, StΓΆhr M | title = Rapid detection of mutagen induced micronucleated erythrocytes by flow cytometry | journal = Histochemistry | volume = 75 | issue = 3 | pages = 353β62 | year = 1982 | pmid = 7141888 | doi = 10.1007/bf00496738 | s2cid = 28973947 | department = Primary }}</ref> Anucleated cells can also arise from flawed cell division in which one daughter lacks a nucleus and the other has two nuclei. In [[flowering plant]]s, this condition occurs in [[sieve tube element]]s.<ref>{{cite journal | vauthors = Ham BK, Lucas WJ | title = The angiosperm phloem sieve tube system: a role in mediating traits important to modern agriculture | journal = Journal of Experimental Botany | volume = 65 | issue = 7 | pages = 1799β816 | date = April 2014 | pmid = 24368503 | doi = 10.1093/jxb/ert417 | doi-access = free }}</ref> ===Multinucleated cells=== {{Main|Multinucleate}} [[Multinucleated]] cells contain multiple nuclei. Most [[acantharea]]n species of [[protozoa]]<ref name="Zettler">{{cite journal | vauthors = Zettler LA, Sogin ML, Caron DA | title = Phylogenetic relationships between the Acantharea and the Polycystinea: a molecular perspective on Haeckel's Radiolaria | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 94 | issue = 21 | pages = 11411β6 | date = October 1997 | pmid = 9326623 | pmc = 23483 | doi = 10.1073/pnas.94.21.11411 | department = Primary | bibcode = 1997PNAS...9411411A | doi-access = free }}</ref> and some [[fungi]] in [[mycorrhizae]]<ref name="Horton">{{cite journal | vauthors = Horton TR | title = The number of nuclei in basidiospores of 63 species of ectomycorrhizal Homobasidiomycetes | journal = Mycologia | volume = 98 | issue = 2 | pages = 233β8 | year = 2006 | pmid = 16894968 | doi = 10.3852/mycologia.98.2.233 | department = Primary }}</ref> have naturally multinucleated cells. Other examples include the [[intestinal parasite]]s in the genus ''[[Giardia]]'', which have two nuclei per cell.<ref>{{cite journal | vauthors = Adam RD | title = The biology of Giardia spp | journal = Microbiological Reviews | volume = 55 | issue = 4 | pages = 706β32 | date = December 1991 | pmid = 1779932 | pmc = 372844 | doi = 10.1128/MMBR.55.4.706-732.1991 | department = Review }}</ref> [[Ciliate]]s have two kinds of nuclei in a single cell, a somatic [[macronucleus]] and a germline [[micronucleus]].<ref>{{cite journal |vauthors=Vogt A, Goldman AD, Mochizuki K, Landweber LF |title=Transposon Domestication versus Mutualism in Ciliate Genome Rearrangements |journal=PLOS Genetics |date=1 August 2013 |volume=9 |issue=8 |pages=e1003659 |doi=10.1371/journal.pgen.1003659 |pmid=23935529 |pmc=3731211 |doi-access=free }}</ref> In humans, [[Skeletal muscle#Skeletal muscle cells|skeletal muscle cells]], also called [[myocyte]]s and [[syncytium]], become multinucleated during development; the resulting arrangement of nuclei near the periphery of the cells allows maximal intracellular space for [[myofibrils]].<ref name="Lodish" /> Other multinucleate cells in the human are [[osteoclast]]s a type of [[bone cell]]. Multinucleated and [[binucleated cells]] can also be abnormal in humans; for example, cells arising from the fusion of [[monocyte]]s and [[macrophage]]s, known as [[giant multinucleated cell]]s, sometimes accompany inflammation<ref name="McInnes">{{cite journal | vauthors = McInnes A, Rennick DM | title = Interleukin 4 induces cultured monocytes/macrophages to form giant multinucleated cells | journal = The Journal of Experimental Medicine | volume = 167 | issue = 2 | pages = 598β611 | date = February 1988 | pmid = 3258008 | pmc = 2188835 | doi = 10.1084/jem.167.2.598 | department = Primary }}</ref> and are also implicated in tumor formation.<ref name="Goldring">{{cite journal | vauthors = Goldring SR, Roelke MS, Petrison KK, Bhan AK | title = Human giant cell tumors of bone identification and characterization of cell types | journal = The Journal of Clinical Investigation | volume = 79 | issue = 2 | pages = 483β91 | date = February 1987 | pmid = 3027126 | pmc = 424109 | doi = 10.1172/JCI112838 | department = Primary }}</ref> A number of [[dinoflagellate]]s are known to have two nuclei. Unlike other multinucleated cells these nuclei contain two distinct lineages of DNA: one from the dinoflagellate and the other from a symbiotic [[diatom]].<ref name=Imanian2012>{{cite journal | vauthors = Imanian B, Pombert JF, Dorrell RG, Burki F, Keeling PJ | title = Tertiary endosymbiosis in two dinotoms has generated little change in the mitochondrial genomes of their dinoflagellate hosts and diatom endosymbionts | journal = PLOS ONE | volume = 7 | issue = 8 | pages = e43763 | year = 2012 | pmid = 22916303 | pmc = 3423374 | doi = 10.1371/journal.pone.0043763 | bibcode = 2012PLoSO...743763I | department = Primary | doi-access = free }}</ref>
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