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Previous studies have demonstrated that the testis-specific SUN domain protein SUN4 (also known as SPAG4) is a vital player in the directed shaping of the spermatid nucleus. However, its molecular properties relating to this crucial function have remained largely unknown, and controversial data for the organization and orientation of SUN4 within the spermatid nuclear envelope have been presented so far. Here, we have re-evaluated this issue in detail and show robust evidence that SUN4 is integral to the inner nuclear membrane, sharing a classical SUN domain protein topology. The C-terminal SUN domain of SUN4 localizes to the perinuclear space, whereas the N-terminus is directed to the nucleoplasm, interacting with the spermiogenesis-specific lamin B3. We found that SUN4 forms heteromeric assemblies with SUN3 in vivo and regulates SUN3 expression. Together, our results contribute to a better understanding of the specific function of SUN4 at the spermatid nucleo-cytoplasmic junction and the process of sperm-head formation.<\/jats:p>","DOI":"10.1242\/jcs.260155","type":"journal-article","created":{"date-parts":[[2023,2,24]],"date-time":"2023-02-24T05:54:36Z","timestamp":1677218076000},"source":"Crossref","is-referenced-by-count":14,"title":["SUN4 is a spermatid type II inner nuclear membrane protein that forms heteromeric assemblies with SUN3 and interacts with lamin B3"],"prefix":"10.1242","volume":"136","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1798-4777","authenticated-orcid":false,"given":"Hanna","family":"Thoma","sequence":"first","affiliation":[{"name":"Biocenter, University of W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"},{"name":"Am Hubland, 97074 W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8924-6589","authenticated-orcid":false,"given":"Luisa","family":"Gr\u00fcnewald","sequence":"additional","affiliation":[{"name":"Biocenter, University of W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"},{"name":"Am Hubland, 97074 W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0196-3453","authenticated-orcid":false,"given":"Silke","family":"Braune","sequence":"additional","affiliation":[{"name":"Biocenter, University of W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"},{"name":"Am Hubland, 97074 W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3183-7861","authenticated-orcid":false,"given":"Elisabeth","family":"Pasch","sequence":"additional","affiliation":[{"name":"Biocenter, University of W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"},{"name":"Am Hubland, 97074 W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6646-556X","authenticated-orcid":false,"given":"Manfred","family":"Alsheimer","sequence":"additional","affiliation":[{"name":"Biocenter, University of W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"},{"name":"Am Hubland, 97074 W\u00fcrzburg Department of Cell and Developmental Biology , , , Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2023,3,27]]},"reference":[{"key":"2024061307541191000_JCS260155C1","doi-asserted-by":"publisher","first-page":"2227","DOI":"10.1242\/jcs.111.15.2227","article-title":"Nuclear envelope remodelling during rat spermiogenesis: distribution and expression pattern of LAP2\/thymopoietins","volume":"111","author":"Alsheimer","year":"1998","journal-title":"J. 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