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In this study, we observed that male knock\u2010out mice for importin \u03b14, which is encoded by the\n                    <jats:italic>Kpna4<\/jats:italic>\n                    gene (\n                    <jats:italic>\n                      Kpna4\n                      <jats:sup>\u2212\/\u2212<\/jats:sup>\n                    <\/jats:italic>\n                    ), were subfertile and yielded smaller litter sizes than those of wild\u2010type (\n                    <jats:italic>WT<\/jats:italic>\n                    ) males. In contrast, mice lacking the closely related importin \u03b13 (\n                    <jats:italic>\n                      Kpna3\n                      <jats:sup>\u2212\/\u2212<\/jats:sup>\n                    <\/jats:italic>\n                    ) were fertile. In vitro fertilization and sperm motility assays demonstrated that sperm from\n                    <jats:italic>\n                      Kpna4\n                      <jats:sup>\u2212\/\u2212<\/jats:sup>\n                    <\/jats:italic>\n                    mice had significantly reduced quality and motility. In addition, acrosome reaction was also impaired in\n                    <jats:italic>\n                      Kpna4\n                      <jats:sup>\u2212\/\u2212<\/jats:sup>\n                    <\/jats:italic>\n                    mice. Transmission electron microscopy revealed striking defects, including abnormal head morphology and multiple axoneme structures in the flagella of\n                    <jats:italic>\n                      Kpna4\n                      <jats:sup>\u2212\/\u2212<\/jats:sup>\n                    <\/jats:italic>\n                    mice. A five\u2010fold increase in the frequency of abnormalities in\n                    <jats:italic>\n                      Kpna4\n                      <jats:sup>\u2212\/\u2212<\/jats:sup>\n                    <\/jats:italic>\n                    mice compared to\n                    <jats:italic>WT<\/jats:italic>\n                    mice indicates the functional importance of importin \u03b14 in normal sperm development. Moreover, Nesprin\u20102, which is a component of the linker of nucleus and cytoskeleton complex, was expressed at lower levels in sperm from\n                    <jats:italic>\n                      Kpna4\n                      <jats:sup>\u2212\/\u2212<\/jats:sup>\n                    <\/jats:italic>\n                    mice and was localized with abnormal axonemes, suggesting incorrect formation of the nuclear membrane\u2010cytoskeleton structure during spermiogenesis. Proteomics analysis of\n                    <jats:italic>\n                      Kpna4\n                      <jats:sup>\u2212\/\u2212<\/jats:sup>\n                    <\/jats:italic>\n                    testis showed significantly altered expression of proteins related to sperm formation, which provided evidence that genetic loss of importin \u03b14 perturbed chromatin status. Collectively, these findings indicate that importin \u03b14 is critical for establishing normal sperm morphology in mice, providing new insights into male germ cell development by highlighting the requirement of importin \u03b14 for normal fertility.\n                  <\/jats:p>","DOI":"10.1096\/fj.202000768rr","type":"journal-article","created":{"date-parts":[[2020,10,15]],"date-time":"2020-10-15T11:30:25Z","timestamp":1602761425000},"page":"16224-16242","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":21,"title":["Genetic loss of importin \u03b14 causes abnormal sperm morphology and impacts on male fertility in mouse"],"prefix":"10.1096","volume":"34","author":[{"given":"Yoichi","family":"Miyamoto","sequence":"first","affiliation":[{"name":"Laboratory of Nuclear Transport Dynamics National Institutes of Biomedical Innovation, Health and Nutrition (NIBIOHN) Osaka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mitsuho","family":"Sasaki","sequence":"additional","affiliation":[{"name":"Laboratory of Animal Models for Human Diseases National Institutes of Biomedical Innovation, Health and Nutrition (NIBIOHN) Osaka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haruhiko","family":"Miyata","sequence":"additional","affiliation":[{"name":"Research Institute for Microbial Diseases Osaka University Osaka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yoko","family":"Monobe","sequence":"additional","affiliation":[{"name":"Section of Laboratory Equipment National Institutes of Biomedical Innovation, Health and Nutrition (NIBIOHN) Osaka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Masahiro","family":"Nagai","sequence":"additional","affiliation":[{"name":"Department of Frontier Bioscience Graduate School of Frontier Bioscience Osaka University Osaka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mayumi","family":"Otani","sequence":"additional","affiliation":[{"name":"Laboratory of Nuclear Transport Dynamics National Institutes of Biomedical Innovation, Health and Nutrition (NIBIOHN) Osaka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Penny A. 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