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<title>Synaptonemal complex</title>

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<span id="openzim-page-title" class="mw-page-title-main"><span class="mw-page-title-main">Synaptonemal complex</span></span>

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<p>The <b>synaptonemal complex</b> (<b>SC</b>) is a <a href="Protein" title="Protein">protein</a> structure that forms between homologous chromosomes (two pairs of <a href="Sister_chromatids" title="Sister chromatids">sister chromatids</a>) during <a href="Meiosis" title="Meiosis">meiosis</a> and is thought to mediate <a href="Synapsis" title="Synapsis">synapsis</a> and <a href="Genetic_recombination" title="Genetic recombination">recombination</a> during prophase I during meiosis in <a href="Eukaryote" title="Eukaryote">eukaryotes</a>. It is currently thought that the SC functions primarily as a scaffold to allow interacting chromatids to complete their <a href="Chromosomal_crossover" title="Chromosomal crossover">crossover</a> activities.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>

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<div class="mw-heading mw-heading2"><h2 id="Composition">Composition</h2></div>

<p>The synaptonemal complex is a tripartite structure consisting of two parallel lateral regions and a central element. This "tripartite structure" is seen during the <a href="Pachytene" title="Pachytene">pachytene</a> stage of the first meiotic <a href="Prophase" title="Prophase">prophase</a>, both in males and in females during <a href="Gametogenesis" title="Gametogenesis">gametogenesis</a>. Previous to the pachytene stage, during leptonema, the lateral elements begin to form and they initiate and complete their pairing during the zygotene stage. After pachynema ends, the SC usually becomes disassembled and can no longer be identified.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>

</p><p>In humans, three specific components of the synaptonemal complex have been characterized: SC protein-1 (SYCP1), SC protein-2 (SYCP2), and SC protein-3 (<a href="SYCP3" title="SYCP3">SYCP3</a>). The SYCP1 <a href="Gene" title="Gene">gene</a> is on chromosome 1p13; the SYCP2 gene is on chromosome 20q13.33; and the gene for SYCP3 is on chromosome 12q.<sup id="cite_ref-:0_3-0" class="reference"><a href="#cite_note-:0-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>

</p><p>The synaptonemal complex was described by Montrose J. Moses in 1956 in primary spermatocytes of crayfish and by D. Fawcett in spermatocytes of pigeon, cat and man.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> As seen with the electron microscope, the synaptonemal complex is formed by two "lateral elements", mainly formed by SYCP3 and secondarily by SYCP2, a "central element" that contains at least two additional proteins and the amino terminal region of SYCP1, and a "central region" spanned between the two lateral elements, that contains the "transverse filaments" composed mainly by the protein SYCP1.<sup id="cite_ref-:0_3-1" class="reference"><a href="#cite_note-:0-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>

</p><p>The SCs can be seen with the light microscope using silver staining or with immunofluorescence techniques that label the proteins SYCP3 or SYCP2.

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<div class="mw-heading mw-heading2"><h2 id="Assembly_and_disassembly">Assembly and disassembly</h2></div>

<p>Formation of the SC usually reflects the pairing or "<a href="Synapsis" title="Synapsis">synapsis</a>" of homologous <a href="Chromosomes" class="mw-redirect" title="Chromosomes">chromosomes</a> and may be used to probe the presence of pairing abnormalities in individuals carrying chromosomal abnormalities, either in number or in the chromosomal structure.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The sex chromosomes in <a href="Male_mammals" class="mw-redirect" title="Male mammals">male mammals</a> show only "partial synapsis" as they usually form only a short SC in the XY pair. The SC shows very little structural variability among eukaryotic organisms despite some significant protein differences. In many organisms the SC carries one or several "recombination nodules" associated with its central space. These nodules are thought to correspond to mature genetic recombination events or "crossovers". In male mice, <a href="Gamma_ray" title="Gamma ray">gamma irradiation</a> increases <a href="Meiosis" title="Meiosis">meiotic crossovers</a> in SCs. This indicates that exogenously caused <a href="DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damages</a> are likely repaired by crossover recombination in SCs.<sup id="cite_ref-:0_3-2" class="reference"><a href="#cite_note-:0-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The finding of an interaction between a SC structural component [synaptonemal central element protein 2 (SYCE2)] and <a href="Homologous_recombination" title="Homologous recombination">recombinational repair</a> protein <a href="RAD51" title="RAD51">RAD51</a> also suggests a role for the SC in DNA repair.

</p><p>In cell development the synaptonemal complex disappears during the late prophase of meiosis I. It is formed during zygotene.

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<div class="mw-heading mw-heading2"><h2 id="Cancer">Cancer</h2></div>

<p>Although synaptonemal complex protein 2 (SYCP2) is a meiotic protein, it is aberrantly and commonly expressed in <a href="Breast_cancer" title="Breast cancer">breast</a> and <a href="Ovarian_cancer" title="Ovarian cancer">ovarian cancers</a>. SYCP2 protein expression in these cancers is associated with broad resistance to drugs that induced <a href="DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damage</a>, i.e. DNA damage response (DDR) drugs.<sup id="cite_ref-Wang2024_6-0" class="reference"><a href="#cite_note-Wang2024-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> SYCP2 is employed in the repair of DNA double-strand breaks by transcription-coupled <a href="Homologous_recombination" title="Homologous recombination">homologous recombination</a>.<sup id="cite_ref-Wang2024_6-1" class="reference"><a href="#cite_note-Wang2024-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> SYCP2 appears to confer cancer cell resistance to therapeutic DNA damaging agents by stimulating R-loop mediated double strand break repair.<sup id="cite_ref-Wang2024_6-2" class="reference"><a href="#cite_note-Wang2024-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Thus inhibition of SYCP2 expression is being studied in efforts to improve therapy for breast and ovarian cancers.<sup id="cite_ref-Wang2024_6-3" class="reference"><a href="#cite_note-Wang2024-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup>

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<div class="mw-heading mw-heading2"><h2 id="Necessity_in_eukaryotes">Necessity in eukaryotes</h2></div>

<p>It is now evident that the synaptonemal complex is not required for genetic recombination in some organisms. For instance, in <a href="Protozoa" title="Protozoa">protozoan</a> <a href="Ciliate" title="Ciliate">ciliates</a> such as <i><a href="Tetrahymena" title="Tetrahymena">Tetrahymena</a> thermophila</i> and <i><a href="Paramecium" title="Paramecium">Paramecium</a> tetraurelia</i> <a href="Chromosomal_crossover" title="Chromosomal crossover">genetic crossover</a> does not appear to require synaptonemal complex formation.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Research has shown that not only does the SC form after genetic recombination but mutant yeast cells unable to assemble a synaptonemal complex can still engage in the exchange of genetic information. However, in other organisms like the <i>C. elegans</i> nematode, formation of chiasmata require the formation of the synaptonemal complex.

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<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>

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</style><cite id="CITEREFPageHawley2004" class="citation journal cs1">Page SL, Hawley RS (2004-10-08). "The genetics and molecular biology of the synaptonemal complex". <i>Annual Review of Cell and Developmental Biology</i>. <b>20</b> (1): <span class="nowrap">525–</span>58. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.cellbio.19.111301.155141">10.1146/annurev.cellbio.19.111301.155141</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/15473851">15473851</a>.</cite></span>

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<li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><cite id="CITEREFYangWang2009" class="citation journal cs1">Yang F, Wang PJ (2009). "The Mammalian synaptonemal complex: a scaffold and beyond". <i>Genome Dynamics</i>. <b>5</b>: <span class="nowrap">69–</span>80. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1159%2F000166620">10.1159/000166620</a>. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-3-8055-8967-3</bdi>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18948708">18948708</a>.</cite></span>

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<li id="cite_note-:0-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-:0_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:0_3-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:0_3-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><cite id="CITEREFBolcun-FilasHallSpeedTaggart2009" class="citation journal cs1">Bolcun-Filas E, Hall E, Speed R, Taggart M, Grey C, de Massy B, et al. (February 2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2640461">"Mutation of the mouse Syce1 gene disrupts synapsis and suggests a link between synaptonemal complex structural components and DNA repair"</a>. <i>PLOS Genetics</i>. <b>5</b> (2): e1000393. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pgen.1000393">10.1371/journal.pgen.1000393</a></span>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2640461">2640461</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19247432">19247432</a>.</cite></span>

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<li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><cite id="CITEREFMoses1968" class="citation journal cs1">Moses, Montrose J. (1968-12-01). "Synaptinemal complex". <i>Annual Review of Genetics</i>. <b>2</b> (1): <span class="nowrap">363–</span>412. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.ge.02.120168.002051">10.1146/annurev.ge.02.120168.002051</a>. <a href="ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0066-4197">0066-4197</a>.</cite></span>

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<li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><cite id="CITEREFZicklerKleckner1999" class="citation journal cs1">Zickler D, Kleckner N (1999-12-01). "Meiotic chromosomes: integrating structure and function". <i>Annual Review of Genetics</i>. <b>33</b> (1): <span class="nowrap">603–</span>754. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.genet.33.1.603">10.1146/annurev.genet.33.1.603</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/10690419">10690419</a>.</cite></span>

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<li id="cite_note-Wang2024-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-Wang2024_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Wang2024_6-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Wang2024_6-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Wang2024_6-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><cite id="CITEREFWangGaoZhangWang2024" class="citation journal cs1">Wang Y, Gao B, Zhang L, Wang X, Zhu X, Yang H, Zhang F, Zhu X, Zhou B, Yao S, Nagayama A, Lee S, Ouyang J, Koh SB, Eisenhauer EL, Zarrella D, Lu K, Rueda BR, Zou L, Su XA, Yeku O, Ellisen LW, Wang XS, Lan L (February 2024). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10881575">"Meiotic protein SYCP2 confers resistance to DNA-damaging agents through R-loop-mediated DNA repair"</a>. <i>Nat Commun</i>. <b>15</b> (1): 1568. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fs41467-024-45693-2">10.1038/s41467-024-45693-2</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10881575">10881575</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/38383600">38383600</a>.</cite><i><span typeof="mw:File"><a href="https://creativecommons.org/licenses/by/4.0/" title="creativecommons:by/4.0/" class="external"></a></span> This article incorporates text from this source, which is available under the <a href="https://creativecommons.org/licenses/by/4.0/" class="extiw external" title="creativecommons:by/4.0/">CC BY 4.0</a> license.</i></span>

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<li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><cite id="CITEREFLukaszewiczHoward-TillLoidl2013" class="citation journal cs1">Lukaszewicz A, Howard-Till RA, Loidl J (November 2013). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3814389">"Mus81 nuclease and Sgs1 helicase are essential for meiotic recombination in a protist lacking a synaptonemal complex"</a>. <i>Nucleic Acids Research</i>. <b>41</b> (20): <span class="nowrap">9296–</span>309. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fnar%2Fgkt703">10.1093/nar/gkt703</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3814389">3814389</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/23935123">23935123</a>.</cite></span>

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<li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text"><cite id="CITEREFChiMahéLoidlLogsdon2014" class="citation journal cs1">Chi J, Mahé F, Loidl J, Logsdon J, Dunthorn M (March 2014). <a rel="nofollow" class="external text" href="https://academic.oup.com/mbe/article/31/3/660/1009587">"Meiosis gene inventory of four ciliates reveals the prevalence of a synaptonemal complex-independent crossover pathway"</a>. <i>Molecular Biology and Evolution</i>. <b>31</b> (3): <span class="nowrap">660–</span>72. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fmolbev%2Fmst258">10.1093/molbev/mst258</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24336924">24336924</a>.</cite></span>

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<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>

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<div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Synaptonemal_complex" class="extiw external" title="commons:Category:Synaptonemal complex">Synaptonemal complex</a></span>.</div></div>

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<ul><li><a rel="nofollow" class="external text" href="http://www.olympusbioscapes.com/gallery/images/333">Synaptonemal complex by 3D-Structured Illumination, photograph by Dr. Chung-Ju Rachel Wang, University of California Berkeley, Department of Molecular and Cell Biology, Berkeley, CA, USA, second place winner of the 2009 Olympus Bioscapes Digital Imaging Competition</a>.</li>

<li>Kounetsova A. et al, <i>Meiosis in Mice without a Synaptonemal Complex</i> PLOS ONE (2011)</li></ul></div><div><div class="zim-footer">

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