{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,11]],"date-time":"2026-05-11T06:10:06Z","timestamp":1778479806975,"version":"3.51.4"},"reference-count":53,"publisher":"Oxford University Press (OUP)","issue":"3","license":[{"start":{"date-parts":[[2025,12,29]],"date-time":"2025-12-29T00:00:00Z","timestamp":1766966400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"DOI":"10.13039\/501100006005","name":"Agence de la Biom\u00e9decine","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100006005","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Inserm and Aix Marseille University"},{"name":"Iraqi Ministry of Higher Education and Scientific Research"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,2,10]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>During spermiogenesis, round spermatids differentiate into spermatozoa. This involves nuclear elongation and condensation, acrosome spreading, and flagellum formation, orchestrated by cytoskeletal elements\u2014acroplaxome and manchette\u2014that attach to the nuclear envelope (NE) except at the caudal pole where nuclear pore complexes (NPCs) form a dense array. Evidence that the NPC contributes structurally to nuclear remodelling comes from a unique case of human male infertility, lacking the spermatid-specific nucleoporin NUP210L, where most sperm heads are round and uncondensed. Mice lacking NUP210L are fertile, however, producing essentially normal spermatozoa. Here, we reveal function at the caudal NE in mice lacking NUP210L and BAF-L, a spermatid-specific paralogue, and interactor, of chromatin protein BAF. Like NUP210L, BAF-L localises at the caudal NE in human elongating spermatids, and its inactivation in mouse has little impact on sperm morphology or fertility. We show here, however, that in mice lacking NUP210L, two copies of BAF-L become essential for fertility. In Banf2+\/\u2212, Nup210l\u2212\/\u2212 and Banf2\u2212\/\u2212, Nup210l\u2212\/\u2212 mice, most spermatids arrest during elongation (step 10\u201311), manchette microtubules frequently invaginate elongating nuclei from the caudal pole, and the NPC array forms at steps 8\u201310 but shows reduced stability later at steps 12\u201316. We show that BAF-L interactor, BAF, also concentrates at the caudal pole (step 8\u20139), independently of NPCs. We propose that BAF-L and NUP210L optimise interactions between NPC, chromatin and the cytoskeleton, to ensure nuclear integrity and fertility. Our study implies that digenic inheritance of mutations in non-homologous genes should be considered when investigating genetic causes of human male infertility.<\/jats:p>","DOI":"10.1093\/hmg\/ddaf194","type":"journal-article","created":{"date-parts":[[2025,12,16]],"date-time":"2025-12-16T12:52:09Z","timestamp":1765889529000},"source":"Crossref","is-referenced-by-count":2,"title":["Nucleoporin NUP210L and BAF-paralogue BAF-L redundantly ensure nuclear integrity and manchette microtubule organisation in mouse spermatids"],"prefix":"10.1093","volume":"35","author":[{"given":"Maha","family":"Al Dala Ali","sequence":"first","affiliation":[{"name":"Aix Marseille Universit\u00e9, Inserm, MMG, U1251 , 27 bd Jean Moulin, 13005 Marseille ,","place":["France"]},{"name":"Al-Iraqia University, College of Medicine, Department of Physiology , Street 22, Mahalla 208, Baghdad ,","place":["Iraq"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Guy","family":"Longepied","sequence":"additional","affiliation":[{"name":"Aix Marseille Universit\u00e9, Inserm, MMG, U1251 , 27 bd Jean Moulin, 13005 Marseille ,","place":["France"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nicolas","family":"L\u00e9vy","sequence":"additional","affiliation":[{"name":"Aix Marseille Universit\u00e9, Inserm, MMG, U1251 , 27 bd Jean Moulin, 13005 Marseille ,","place":["France"]},{"name":"Servier , Research and Development, 91190 Gif-sur-Yvette,","place":["France"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Catherine","family":"Metzler-Guillemain","sequence":"additional","affiliation":[{"name":"Aix Marseille Universit\u00e9, Inserm, MMG, U1251 , 27 bd Jean Moulin, 13005 Marseille ,","place":["France"]},{"name":"APHM, H\u00f4pital de la Conception, P\u00f4le Femmes-Parents-Enfants, Centre Clinico-biologique AMP-CECOS , 147 bd Baille, 13005 Marseille ,","place":["France"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2052-7347","authenticated-orcid":false,"given":"Michael J","family":"Mitchell","sequence":"additional","affiliation":[{"name":"Aix Marseille Universit\u00e9, Inserm, MMG, U1251 , 27 bd Jean Moulin, 13005 Marseille ,","place":["France"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2025,12,29]]},"reference":[{"key":"2026051101331573400_ref1","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1679\/aohc.67.271","article-title":"The acrosome-acroplaxome-manchette complex and the shaping of the spermatid head","volume":"67","author":"Kierszenbaum","year":"2004","journal-title":"Arch Histol Cytol"},{"key":"2026051101331573400_ref2","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1016\/j.ydbio.2015.09.010","article-title":"SUN4 is essential for nuclear remodeling during 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