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FANCD2
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<h2 id="function">Function</h2> <p><a href="/facts/Fanconi_anemia/4p1QUprP">Fanconi anemia</a> is a disorder with a recessive Mendelian pattern of inheritance characterized by <a href="/facts/Chromosome_instability_syndrome/F2ubfxkY">chromosomal instability</a>, hypersensitivity to DNA crosslinking agents, increased <a href="/facts/DNA_repair/hsURuYYq">chromosomal breakage</a>, and defective <a href="/facts/DNA_repair/hsURuYYq">DNA repair</a>. The members of the Fanconi anemia complementation group do not share sequence similarity; they are related by their assembly into a common nuclear protein complex. This gene encodes the protein for complementation group D2. This protein is <a href="/facts/Ubiquitin/hpulmU3m">monoubiquitinated</a> in response to DNA damage, resulting in its localization to nuclear foci with other proteins (<a href="/facts/BRCA1/xHDwxARA">BRCA1</a> and <a href="/facts/BRCA2/eDUI83G1">BRCA2</a>) involved in homology-directed DNA repair (see Figure: Recombinational repair of DNA double-strand damages). A <a href="/facts/Cell_nucleus/D7sFRa3c">nuclear</a> complex containing <a href="/facts/FANCA/9RdnPrbG">FANCA</a>, [Fanconi anemia, complementation group A], <a href="/facts/FANCB/qRfKdFmc">FANCB</a>, <a href="/facts/FANCC/j3dGjnYe">FANCC</a>, <a href="/facts/FANCE/IMtHx2xp">FANCE</a>, <a href="/facts/FANCF/PHGt9fss">FANCF</a>, <a href="/facts/FANCL/NneVA3k8">FANCL</a> and <a href="/facts/FANCG/dd8SvMsk">FANCG</a> proteins is required for the activation of the FANCD2 protein to the <a href="/facts/Ubiquitin/hpulmU3m">mono-ubiquitinated</a> isoform.<a class="footnote-ref" id="fnref:4" href="#fn:4"><sup>4</sup></a> </p><p>Mono-ubiquination of FANCD2 is essential for repairing DNA interstrand crosslinks, and clamps the protein on DNA together with its partner protein <a href="/facts/FANCI/RqQTvSBC">FANCI</a>. The monoubiquitinated FANCD2:FANCI complex coats DNA in a filament-like array, potentially as a way to protect DNA associated with stalled replication.<a class="footnote-ref" id="fnref:5" href="#fn:5"><sup>5</sup></a> </p><p>Mono-ubiquitination is also required for interaction with the nuclease <a href="/facts/FAN1/dlRVUoto">FAN1</a>. <a href="/facts/FAN1/dlRVUoto">FAN1</a> recruitment and its consequent activity restrain DNA replication fork progression and prevent chromosome abnormalities from occurring when DNA replication forks stall.<a class="footnote-ref" id="fnref:6" href="#fn:6"><sup>6</sup></a> </p> <h2 id="infertility">Infertility</h2> <p>Humans with a FANCD deficiency display hypogonadism, male infertility, impaired spermatogenesis, and reduced female fertility. Similarly, mice deficient in FANCD2 show hypogonadism, impaired fertility and impaired gametogenesis.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> </p><p>In the non-mutant mouse, FANCD2 is expressed in <a href="/facts/Spermatogonium/wIb48yRh">spermatogonia</a>, pre-leptotene spermatocytes, and in spermatocytes in the leptotene, zygotene and early pachytene stages of <a href="/facts/Meiosis/Pj5Q6nGN">meiosis</a>.<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a> In <a href="/facts/Synaptonemal_complex/vTadctRx">synaptonemal complexes</a> of meiotic chromosomes, activated FANCD2 protein co-localizes with <a href="/facts/BRCA1/xHDwxARA">BRCA1</a> (breast cancer susceptibility protein).<a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a> FANCD2 mutant mice exhibit chromosome mis-pairing during the pachytene stage of meiosis and <a href="/facts/Germ_cell/1kY5KaR1">germ cell</a> loss.<a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a> Activated FANCD2 protein may normally function prior to the initiation of meiotic recombination, perhaps to prepare chromosomes for synapsis, or to regulate subsequent recombination events.<a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a> </p> <h2 id="clinical-significance">Clinical significance</h2> <p>Tobacco smoke suppresses the expression of FANCD2, which codes for a DNA damage "caretaker" or repair mechanism.<a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a> </p> <h3>Cancer</h3> <p>FANCD2 mutant mice have a significantly increased incidence of tumors including ovarian, gastric and hepatic adenomas as well as hepatocellular, lung, ovarian and mammary carcinomas.<a class="footnote-ref" id="fnref:13" href="#fn:13"><sup>13</sup></a><a class="footnote-ref" id="fnref:14" href="#fn:14"><sup>14</sup></a> Humans with a FANCD2 deficiency have increased acute myeloid leukemia, and squamous cell carcinomas (head and neck squamous cell carcinomas and anogenital carcinomas).<a class="footnote-ref" id="fnref:15" href="#fn:15"><sup>15</sup></a> Lung squamous tumors express high levels of FANCD2 and members of Fanconia anemia pathway.<a class="footnote-ref" id="fnref:16" href="#fn:16"><sup>16</sup></a> </p><p>FANCD2 monoubiquitination is also a potential therapeutic target in the treatment of cancer.<a class="footnote-ref" id="fnref:17" href="#fn:17"><sup>17</sup></a> </p> <h2 id="interactions">Interactions</h2> <p>FANCD2 has been shown to <a href="/facts/Protein-protein_interaction/etvm9Wmg">interact</a> with: </p> <ul><li><a href="/facts/FANCI/RqQTvSBC">FANCI</a><a class="footnote-ref" id="fnref:18" href="#fn:18"><sup>18</sup></a><a class="footnote-ref" id="fnref:19" href="#fn:19"><sup>19</sup></a></li> <li><a href="/facts/Ataxia_telangiectasia_mutated/oBMxx2Y7">Ataxia telangiectasia mutated</a>,<a class="footnote-ref" id="fnref:20" href="#fn:20"><sup>20</sup></a><a class="footnote-ref" id="fnref:21" href="#fn:21"><sup>21</sup></a></li> <li><a href="/facts/BARD1/9yN8kBh5">BARD1</a>,<a class="footnote-ref" id="fnref:22" href="#fn:22"><sup>22</sup></a></li> <li><a href="/facts/BRCA1/xHDwxARA">BRCA1</a>.<a class="footnote-ref" id="fnref:23" href="#fn:23"><sup>23</sup></a><a class="footnote-ref" id="fnref:24" href="#fn:24"><sup>24</sup></a></li> <li><a href="/facts/BRCA2/eDUI83G1">BRCA2</a>,<a class="footnote-ref" id="fnref:25" href="#fn:25"><sup>25</sup></a><a class="footnote-ref" id="fnref:26" href="#fn:26"><sup>26</sup></a><a class="footnote-ref" id="fnref:27" href="#fn:27"><sup>27</sup></a></li> <li><a href="/facts/FANCE/IMtHx2xp">FANCE</a>,<a class="footnote-ref" id="fnref:28" href="#fn:28"><sup>28</sup></a><a class="footnote-ref" id="fnref:29" href="#fn:29"><sup>29</sup></a><a class="footnote-ref" id="fnref:30" href="#fn:30"><sup>30</sup></a></li> <li><a href="/facts/HTATIP/UkyldBkS">HTATIP</a>,<a class="footnote-ref" id="fnref:31" href="#fn:31"><sup>31</sup></a> and</li> <li><a href="/facts/MEN1/SNDm7wqX">MEN1</a>.<a class="footnote-ref" id="fnref:32" href="#fn:32"><sup>32</sup></a></li></ul> <h2 id="external-links">External links</h2> <h2 id="further-reading">Further reading</h2> <ul><li>Hejna JA, Timmers CD, Reifsteck C, Bruun DA, Lucas LW, Jakobs PM, et al. (May 2000). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1378015">"Localization of the Fanconi anemia complementation group D gene to a 200-kb region on chromosome 3p25.3"</a>. <i>American Journal of Human Genetics</i>. 66 (5): 1540–51. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1086%2F302896">10.1086/302896</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1378015">1378015</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/10762542">10762542</a>.</li> <li>Garcia-Higuera I, Taniguchi T, Ganesan S, Meyn MS, Timmers C, Hejna J, et al. (February 2001). <a href="https://doi.org/10.1016%2FS1097-2765%2801%2900173-3">"Interaction of the Fanconi anemia proteins and BRCA1 in a common pathway"</a>. <i>Molecular Cell</i>. 7 (2): 249–62. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2FS1097-2765%2801%2900173-3">10.1016/S1097-2765(01)00173-3</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/11239454">11239454</a>.</li> <li>Futaki M, Liu JM (December 2001). "Chromosomal breakage syndromes and the BRCA1 genome surveillance complex". <i>Trends in Molecular Medicine</i>. 7 (12): 560–5. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2FS1471-4914%2801%2902178-5">10.1016/S1471-4914(01)02178-5</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/11733219">11733219</a>.</li> <li>Wilson JB, Johnson MA, Stuckert AP, Trueman KL, May S, Bryant PE, et al. (December 2001). <a href="https://doi.org/10.1093%2Fcarcin%2F22.12.1939">"The Chinese hamster FANCG/XRCC9 mutant NM3 fails to express the monoubiquitinated form of the FANCD2 protein, is hypersensitive to a range of DNA damaging agents and exhibits a normal level of spontaneous sister chromatid exchange"</a>. <i>Carcinogenesis</i>. 22 (12): 1939–46. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1093%2Fcarcin%2F22.12.1939">10.1093/carcin/22.12.1939</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/11751423">11751423</a>.</li> <li>Grompe M (June 2002). "FANCD2: a branch-point in DNA damage response?". <i>Nature Medicine</i>. 8 (6): 555–6. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1038%2Fnm0602-555">10.1038/nm0602-555</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12042798">12042798</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:27843912">27843912</a>.</li> <li>Taniguchi T, Garcia-Higuera I, Xu B, Andreassen PR, Gregory RC, Kim ST, et al. (May 2002). <a href="https://doi.org/10.1016%2FS0092-8674%2802%2900747-X">"Convergence of the fanconi anemia and ataxia telangiectasia signaling pathways"</a>. <i>Cell</i>. 109 (4): 459–72. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2FS0092-8674%2802%2900747-X">10.1016/S0092-8674(02)00747-X</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12086603">12086603</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:16580666">16580666</a>.</li> <li>Pace P, Johnson M, Tan WM, Mosedale G, Sng C, Hoatlin M, et al. (July 2002). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC125396">"FANCE: the link between Fanconi anaemia complex assembly and activity"</a>. <i>The EMBO Journal</i>. 21 (13): 3414–23. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1093%2Femboj%2Fcdf355">10.1093/emboj/cdf355</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC125396">125396</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12093742">12093742</a>.</li> <li>Taniguchi T, Garcia-Higuera I, Andreassen PR, Gregory RC, Grompe M, D'Andrea AD (October 2002). <a href="https://doi.org/10.1182%2Fblood-2002-01-0278">"S-phase-specific interaction of the Fanconi anemia protein, FANCD2, with BRCA1 and RAD51"</a>. <i>Blood</i>. 100 (7): 2414–20. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1182%2Fblood-2002-01-0278">10.1182/blood-2002-01-0278</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12239151">12239151</a>.</li> <li>Tamary H, Bar-Yam R, Zemach M, Dgany O, Shalmon L, Yaniv I (October 2002). "The molecular biology of Fanconi anemia". <i>The Israel Medical Association Journal</i>. 4 (10): 819–23. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12389351">12389351</a>.</li> <li>Nakanishi K, Taniguchi T, Ranganathan V, New HV, Moreau LA, Stotsky M, et al. (December 2002). "Interaction of FANCD2 and NBS1 in the DNA damage response". <i>Nature Cell Biology</i>. 4 (12): 913–20. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1038%2Fncb879">10.1038/ncb879</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12447395">12447395</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:20807784">20807784</a>.</li> <li>Goldberg M, Stucki M, Falck J, D'Amours D, Rahman D, Pappin D, et al. (February 2003). "MDC1 is required for the intra-S-phase DNA damage checkpoint". <i>Nature</i>. 421 (6926): 952–6. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/2003Natur.421..952G">2003Natur.421..952G</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1038%2Fnature01445">10.1038/nature01445</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12607003">12607003</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:4301037">4301037</a>.</li> <li>Stewart GS, Wang B, Bignell CR, Taylor AM, Elledge SJ (February 2003). "MDC1 is a mediator of the mammalian DNA damage checkpoint". <i>Nature</i>. 421 (6926): 961–6. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/2003Natur.421..961S">2003Natur.421..961S</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1038%2Fnature01446">10.1038/nature01446</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12607005">12607005</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:4410773">4410773</a>.</li> <li>Gordon SM, Buchwald M (July 2003). "Fanconi anemia protein complex: mapping protein interactions in the yeast 2- and 3-hybrid systems". <i>Blood</i>. 102 (1): 136–41. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1182%2Fblood-2002-11-3517">10.1182/blood-2002-11-3517</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12649160">12649160</a>.</li> <li>Jin S, Mao H, Schnepp RW, Sykes SM, Silva AC, D'Andrea AD, Hua X (July 2003). "Menin associates with FANCD2, a protein involved in repair of DNA damage". <i>Cancer Research</i>. 63 (14): 4204–10. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12874027">12874027</a>.</li> <li>Vandenberg CJ, Gergely F, Ong CY, Pace P, Mallery DL, Hiom K, Patel KJ (July 2003). <a href="https://doi.org/10.1016%2FS1097-2765%2803%2900281-8">"BRCA1-independent ubiquitination of FANCD2"</a>. <i>Molecular Cell</i>. 12 (1): 247–54. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2FS1097-2765%2803%2900281-8">10.1016/S1097-2765(03)00281-8</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12887909">12887909</a>.</li> <li>Meetei AR, de Winter JP, Medhurst AL, Wallisch M, Waisfisz Q, van de Vrugt HJ, et al. (October 2003). "A novel ubiquitin ligase is deficient in Fanconi anemia". <i>Nature Genetics</i>. 35 (2): 165–70. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1038%2Fng1241">10.1038/ng1241</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/12973351">12973351</a>. <a href="/facts/S2CID_(identifier)/ldJsHa2Y">S2CID</a> <a href="https://api.semanticscholar.org/CorpusID:10149290">10149290</a>.</li> <li>Reuter TY, Medhurst AL, Waisfisz Q, Zhi Y, Herterich S, Hoehn H, et al. (October 2003). "Yeast two-hybrid screens imply involvement of Fanconi anemia proteins in transcription regulation, cell signaling, oxidative metabolism, and cellular transport". <i>Experimental Cell Research</i>. 289 (2): 211–21. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2FS0014-4827%2803%2900261-1">10.1016/S0014-4827(03)00261-1</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/14499622">14499622</a>.</li> <li>Wang X, Kennedy RD, Ray K, Stuckert P, Ellenberger T, D'Andrea AD (April 2007). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1899922">"Chk1-mediated phosphorylation of FANCE is required for the Fanconi anemia/BRCA pathway"</a>. <i>Molecular and Cellular Biology</i>. 27 (8): 3098–108. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1128%2FMCB.02357-06">10.1128/MCB.02357-06</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1899922">1899922</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/17296736">17296736</a>.</li></ul> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Whitney M, Thayer M, Reifsteck C, Olson S, Smith L, Jakobs PM, et al. (November 1995). "Microcell mediated chromosome transfer maps the Fanconi anaemia group D gene to chromosome 3p". Nature Genetics. 11 (3): 341–3. doi:10.1038/ng1195-341. PMID 7581463. S2CID 451042. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Timmers C, Taniguchi T, Hejna J, Reifsteck C, Lucas L, Bruun D, et al. (February 2001). "Positional cloning of a novel Fanconi anemia gene, FANCD2". Molecular Cell. 7 (2): 241–8. doi:10.1016/S1097-2765(01)00172-1. PMID 11239453. <a href="https://doi.org/10.1016%2FS1097-2765%2801%2900172-1" target="_blank">https://doi.org/10.1016%2FS1097-2765%2801%2900172-1</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>Sharp, Michael F.; Bythell-Douglas, Rohan; Deans, Andrew J.; Crismani, Wayne (2021). "The Fanconi anemia ubiquitin E3 ligase complex as an anti-cancer target". Molecular Cell. 81 (11): 2278–2289. doi:10.1016/j.molcel.2021.04.023. <a href="https://linkinghub.elsevier.com/retrieve/pii/S1097276521003282" target="_blank">https://linkinghub.elsevier.com/retrieve/pii/S1097276521003282</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>Garcia-Higuera I, Taniguchi T, Ganesan S, Meyn MS, Timmers C, Hejna J, et al. (February 2001). "Interaction of the Fanconi anemia proteins and BRCA1 in a common pathway". Molecular Cell. 7 (2): 249–62. doi:10.1016/s1097-2765(01)00173-3. PMID 11239454. <a href="https://doi.org/10.1016%2Fs1097-2765%2801%2900173-3" target="_blank">https://doi.org/10.1016%2Fs1097-2765%2801%2900173-3</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>Tan W, van Twest S, Leis A, Bythell-Douglas R, Murphy VJ, Sharp M, et al. (March 2020). 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