{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,25]],"date-time":"2026-07-25T07:48:31Z","timestamp":1784965711754,"version":"3.55.0"},"reference-count":34,"publisher":"Springer Science and Business Media LLC","issue":"5","license":[{"start":{"date-parts":[[2021,6,2]],"date-time":"2021-06-02T00:00:00Z","timestamp":1622592000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"},{"start":{"date-parts":[[2021,6,2]],"date-time":"2021-06-02T00:00:00Z","timestamp":1622592000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"}],"funder":[{"name":"the National Key Research and Developmental Program of China","award":["2018YFC1003403 and 2016YFC1000600"],"award-info":[{"award-number":["2018YFC1003403 and 2016YFC1000600"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["31890780 and 31630050"],"award-info":[{"award-number":["31890780 and 31630050"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Strategic Priority Research Program of the Chinese Academy of Sciences","award":["XDB19000000"],"award-info":[{"award-number":["XDB19000000"]}]},{"DOI":"10.13039\/501100012226","name":"Fundamental Research Funds for the Central Universities","doi-asserted-by":"publisher","award":["YD2070002006"],"award-info":[{"award-number":["YD2070002006"]}],"id":[{"id":"10.13039\/501100012226","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Mamm Genome"],"published-print":{"date-parts":[[2021,10]]},"DOI":"10.1007\/s00335-021-09879-z","type":"journal-article","created":{"date-parts":[[2021,6,2]],"date-time":"2021-06-02T08:03:06Z","timestamp":1622620986000},"page":"364-370","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":8,"title":["Inactivation of testis-specific gene C4orf46 is dispensable for spermatogenesis and fertility in mouse"],"prefix":"10.1007","volume":"32","author":[{"given":"Basit","family":"Shah","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1010-4407","authenticated-orcid":false,"given":"Ranjha","family":"Khan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wasim","family":"Shah","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ayesha","family":"Aftab","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Manan","family":"Khan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sobia","family":"Dil","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qinghua","family":"Shi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2021,6,2]]},"reference":[{"issue":"15","key":"9879_CR1","doi-asserted-by":"publisher","first-page":"6346","DOI":"10.1128\/MCB.25.15.6346-6354.2005","volume":"25","author":"N Bedard","year":"2005","unstructured":"Bedard N et al (2005) Mice lacking the UBC4-testis gene have a delay in postnatal testis development but normal spermatogenesis and fertility. Mol Cell Biol 25(15):6346\u20136354","journal-title":"Mol Cell Biol"},{"issue":"10","key":"9879_CR2","doi-asserted-by":"publisher","first-page":"1429","DOI":"10.1038\/cr.2012.120","volume":"22","author":"E Beyret","year":"2012","unstructured":"Beyret E, Liu N, Lin H (2012) piRNA biogenesis during adult spermatogenesis in mice is independent of the ping-pong mechanism. Cell Res 22(10):1429\u20131439","journal-title":"Cell Res"},{"issue":"27","key":"9879_CR3","doi-asserted-by":"publisher","first-page":"E5370","DOI":"10.1073\/pnas.1621279114","volume":"114","author":"JM Castaneda","year":"2017","unstructured":"Castaneda JM et al (2017) TCTE1 is a conserved component of the dynein regulatory complex and is required for motility and metabolism in mouse spermatozoa. Proc Natl Acad Sci USA 114(27):E5370\u2013E5378","journal-title":"Proc Natl Acad Sci USA"},{"issue":"3","key":"9879_CR4","doi-asserted-by":"publisher","first-page":"209","DOI":"10.4161\/spmg.1.3.17990","volume":"1","author":"YH Cheng","year":"2011","unstructured":"Cheng YH, Wong EW, Cheng CY (2011) Cancer\/testis (CT) antigens, carcinogenesis and spermatogenesis. Spermatogenesis 1(3):209\u2013220","journal-title":"Spermatogenesis"},{"issue":"1","key":"9879_CR5","doi-asserted-by":"publisher","first-page":"017","DOI":"10.1038\/s41467-017-00019-3","volume":"8","author":"D Ding","year":"2017","unstructured":"Ding D et al (2017) PNLDC1 is essential for piRNA 3\u2019 end trimming and transposon silencing during spermatogenesis in mice. Nat Commun 8(1):017\u201300854","journal-title":"Nat Commun"},{"issue":"19","key":"9879_CR6","doi-asserted-by":"publisher","first-page":"6289","DOI":"10.1074\/jbc.RA119.012375","volume":"295","author":"Q Gao","year":"2020","unstructured":"Gao Q et al (2020) The testis-specific LINC component SUN3 is essential for sperm head shaping during mouse spermiogenesis. J Biol Chem 295(19):6289\u20136298","journal-title":"J Biol Chem"},{"issue":"1","key":"9879_CR7","first-page":"84","volume":"102","author":"RJ Holcomb","year":"2020","unstructured":"Holcomb RJ et al (2020) The testis-specific serine proteases PRSS44, PRSS46, and PRSS54 are dispensable for male mouse fertility\u2020. Biol Reprod 102(1):84\u201391","journal-title":"Biol Reprod"},{"issue":"22","key":"9879_CR8","first-page":"11755","volume":"47","author":"R Hua","year":"2019","unstructured":"Hua R et al (2019) FBXO47 regulates telomere-inner nuclear envelope integration by stabilizing TRF2 during meiosis. Nucleic Acids Res 47(22):11755\u201311770","journal-title":"Nucleic Acids Res"},{"issue":"4","key":"9879_CR9","doi-asserted-by":"publisher","first-page":"544","DOI":"10.1007\/s11427-018-9469-4","volume":"62","author":"Z Huang","year":"2019","unstructured":"Huang Z et al (2019) The deubiquitinating gene Usp29 is dispensable for fertility in male mice. Sci China Life Sci 62(4):544\u2013552","journal-title":"Sci China Life Sci"},{"key":"9879_CR10","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/15548627.2020.1816342","volume":"17","author":"Q Huang","year":"2020","unstructured":"Huang Q et al (2020) Autophagy core protein ATG5 is required for elongating spermatid development, sperm individualization and normal fertility in male mice. Autophagy 17:1\u201315","journal-title":"Autophagy"},{"issue":"6","key":"9879_CR11","doi-asserted-by":"publisher","first-page":"1225","DOI":"10.1095\/biolreprod.110.088955","volume":"84","author":"N Iwamori","year":"2011","unstructured":"Iwamori N et al (2011) The testis-enriched histone demethylase, KDM4D, regulates methylation of histone H3 lysine 9 during spermatogenesis in the mouse but is dispensable for fertility. Biol Reprod 84(6):1225\u20131234","journal-title":"Biol Reprod"},{"issue":"10","key":"9879_CR12","doi-asserted-by":"publisher","first-page":"1498","DOI":"10.1016\/j.cub.2017.04.017","volume":"27","author":"L Jiang","year":"2017","unstructured":"Jiang L et al (2017) RPL10L Is required for male meiotic division by compensating for RPL10 during meiotic sex chromosome inactivation in mice. Curr Biol 27(10):1498\u20131505","journal-title":"Curr Biol"},{"issue":"1","key":"9879_CR13","doi-asserted-by":"publisher","first-page":"018","DOI":"10.1038\/s41598-017-18535-z","volume":"8","author":"M Khan","year":"2018","unstructured":"Khan M et al (2018) The evolutionarily conserved genes: Tex37, Ccdc73, Prss55 and Nxt2 are dispensable for fertility in mice. Sci Rep 8(1):018\u201323176","journal-title":"Sci Rep"},{"issue":"7","key":"9879_CR14","doi-asserted-by":"publisher","first-page":"5207","DOI":"10.1007\/s11033-020-05595-0","volume":"47","author":"R Khan","year":"2020","unstructured":"Khan R et al (2020) Evolutionarily conserved and testis-specific gene, 4930524B15Rik, is not essential for mouse spermatogenesis and fertility. Mol Biol Rep 47(7):5207\u20135213","journal-title":"Mol Biol Rep"},{"issue":"9","key":"9879_CR15","doi-asserted-by":"publisher","first-page":"809","DOI":"10.1016\/j.tree.2020.04.009","volume":"35","author":"\u00c1 L\u00e1ruson","year":"2020","unstructured":"L\u00e1ruson \u00c1, Yeaman JS, Lotterhos KE (2020) The importance of genetic redundancy in evolution. Trends Ecol Evol 35(9):809\u2013822","journal-title":"Trends Ecol Evol"},{"issue":"6","key":"9879_CR16","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1093\/molehr\/gau014","volume":"20","author":"M Liu","year":"2014","unstructured":"Liu M et al (2014) SHCBP1L, a conserved protein in mammals, is predominantly expressed in male germ cells and maintains spindle stability during meiosis in testis. Mol Hum Reprod 20(6):463\u2013475","journal-title":"Mol Hum Reprod"},{"issue":"2","key":"9879_CR17","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1093\/biolre\/ioz103","volume":"101","author":"Y Lu","year":"2019","unstructured":"Lu Y et al (2019) CRISPR\/Cas9-mediated genome editing reveals 30 testis-enriched genes dispensable for male fertility in mice\u2020. Biol Reprod 101(2):501\u2013511","journal-title":"Biol Reprod"},{"issue":"1","key":"9879_CR18","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1016\/j.ceca.2014.10.009","volume":"58","author":"MR Miller","year":"2015","unstructured":"Miller MR et al (2015) Flagellar ion channels of sperm: similarities and differences between species. Cell Calcium 58(1):105\u2013113","journal-title":"Cell Calcium"},{"issue":"12","key":"9879_CR19","doi-asserted-by":"publisher","first-page":"16224","DOI":"10.1096\/fj.202000768RR","volume":"34","author":"Y Miyamoto","year":"2020","unstructured":"Miyamoto Y et al (2020) Genetic loss of importin \u03b14 causes abnormal sperm morphology and impacts on male fertility in mouse. Faseb J 34(12):16224\u201316242","journal-title":"Faseb J"},{"issue":"28","key":"9879_CR20","doi-asserted-by":"publisher","first-page":"7704","DOI":"10.1073\/pnas.1608458113","volume":"113","author":"H Miyata","year":"2016","unstructured":"Miyata H et al (2016) Genome engineering uncovers 54 evolutionarily conserved and testis-enriched genes that are not required for male fertility in mice. Proc Natl Acad Sci USA 113(28):7704\u20137710","journal-title":"Proc Natl Acad Sci USA"},{"issue":"1","key":"9879_CR21","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1007\/s11033-019-05131-9","volume":"47","author":"D Nie","year":"2020","unstructured":"Nie D, Dai Y, Luo Z (2020) The testis-specific expressed gene Spata34 is not required for fertility in mice. Mol Biol Rep 47(1):285\u2013292","journal-title":"Mol Biol Rep"},{"issue":"5","key":"9879_CR22","doi-asserted-by":"publisher","first-page":"390","DOI":"10.1002\/mrd.22319","volume":"81","author":"S Ozturk","year":"2014","unstructured":"Ozturk S et al (2014) Epab is dispensable for mouse spermatogenesis and male fertility. Mol Reprod Dev 81(5):390. https:\/\/doi.org\/10.1002\/mrd.22319","journal-title":"Mol Reprod Dev"},{"issue":"4","key":"9879_CR23","doi-asserted-by":"publisher","first-page":"1219","DOI":"10.1073\/pnas.0610286104","volume":"104","author":"H Qi","year":"2007","unstructured":"Qi H et al (2007) All four CatSper ion channel proteins are required for male fertility and sperm cell hyperactivated motility. Proc Natl Acad Sci USA 104(4):1219\u20131223","journal-title":"Proc Natl Acad Sci USA"},{"key":"9879_CR24","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1034\/j.1600-065X.2002.18803.x","volume":"188","author":"MJ Scanlan","year":"2002","unstructured":"Scanlan MJ et al (2002) Cancer\/testis antigens: an expanding family of targets for cancer immunotherapy. Immunol Rev 188:22\u201332","journal-title":"Immunol Rev"},{"issue":"21","key":"9879_CR25","doi-asserted-by":"publisher","first-page":"12201","DOI":"10.1073\/pnas.1635054100","volume":"100","author":"N Schultz","year":"2003","unstructured":"Schultz N, Hamra FK, Garbers DL (2003) A multitude of genes expressed solely in meiotic or postmeiotic spermatogenic cells offers a myriad of contraceptive targets. Proc Natl Acad Sci USA 100(21):12201\u201312206","journal-title":"Proc Natl Acad Sci USA"},{"issue":"8","key":"9879_CR27","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1038\/nrc1669","volume":"5","author":"AJ Simpson","year":"2005","unstructured":"Simpson AJ et al (2005) Cancer\/testis antigens, gametogenesis and cancer. Nat Rev Cancer 5(8):615\u2013625","journal-title":"Nat Rev Cancer"},{"issue":"22","key":"9879_CR28","doi-asserted-by":"publisher","first-page":"9325","DOI":"10.1073\/pnas.0611296104","volume":"104","author":"D Wang","year":"2007","unstructured":"Wang D et al (2007) A sperm-specific Na+\/H+ exchanger (sNHE) is critical for expression and in vivo bicarbonate regulation of the soluble adenylyl cyclase (sAC). Proc Natl Acad Sci USA 104(22):9325\u20139330","journal-title":"Proc Natl Acad Sci USA"},{"issue":"143925","key":"9879_CR29","first-page":"15","volume":"711","author":"Y Xie","year":"2019","unstructured":"Xie Y et al (2019) The testis-specifically expressed Dpep3 is not essential for male fertility in mice. Gene 711(143925):15","journal-title":"Gene"},{"issue":"4","key":"9879_CR30","doi-asserted-by":"publisher","first-page":"520","DOI":"10.1016\/j.repbio.2020.08.006","volume":"20","author":"A Yousaf","year":"2020","unstructured":"Yousaf A et al (2020) Normal spermatogenesis and fertility in Ddi1 (DNA damage inducible 1) mutant mice. Reprod Biol 20(4):520\u2013524","journal-title":"Reprod Biol"},{"issue":"3","key":"9879_CR31","doi-asserted-by":"publisher","first-page":"1583","DOI":"10.3892\/mmr.2014.2388","volume":"10","author":"Z Yu","year":"2014","unstructured":"Yu Z et al (2014) Expression and clinical significance of RCDG1 in renal cell carcinoma: a novel renal cancer-associated gene. Mol Med Rep 10(3):1583\u20131589","journal-title":"Mol Med Rep"},{"issue":"145254","key":"9879_CR32","first-page":"22","volume":"769","author":"Q Zaman","year":"2021","unstructured":"Zaman Q et al (2021) Exonuclease 5 is dispensable for meiotic progression and male fertility in mouse. Gene 769(145254):22","journal-title":"Gene"},{"issue":"3","key":"9879_CR33","doi-asserted-by":"publisher","first-page":"272","DOI":"10.1002\/jcp.10215","volume":"194","author":"AJ Zendman","year":"2003","unstructured":"Zendman AJ, Ruiter DJ, Van Muijen GN (2003) Cancer\/testis-associated genes: identification, expression profile, and putative function. J Cell Physiol 194(3):272\u2013288","journal-title":"J Cell Physiol"},{"issue":"14","key":"9879_CR34","doi-asserted-by":"publisher","first-page":"5879","DOI":"10.1073\/pnas.1100240108","volume":"108","author":"XH Zeng","year":"2011","unstructured":"Zeng XH et al (2011) Deletion of the Slo3 gene abolishes alkalization-activated K+ current in mouse spermatozoa. Proc Natl Acad Sci USA 108(14):5879\u20135884","journal-title":"Proc Natl Acad Sci USA"},{"key":"9879_CR35","doi-asserted-by":"publisher","first-page":"e6827","DOI":"10.7717\/peerj.6827","volume":"7","author":"J Zhang","year":"2019","unstructured":"Zhang J et al (2019) Normal spermatogenesis in Fank1 (fibronectin type 3 and ankyrin repeat domains 1) mutant mice. Peer J 7:e6827","journal-title":"Peer J"}],"container-title":["Mammalian Genome"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00335-021-09879-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00335-021-09879-z\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00335-021-09879-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,9,22]],"date-time":"2021-09-22T12:03:54Z","timestamp":1632312234000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00335-021-09879-z"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,6,2]]},"references-count":34,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2021,10]]}},"alternative-id":["9879"],"URL":"https:\/\/doi.org\/10.1007\/s00335-021-09879-z","relation":{},"ISSN":["0938-8990","1432-1777"],"issn-type":[{"value":"0938-8990","type":"print"},{"value":"1432-1777","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,6,2]]},"assertion":[{"value":"6 March 2021","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 May 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 June 2021","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"All authors declare that no conflict of interest exists.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Conflict of interest"}},{"value":"Included in materials and methods section.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval"}},{"value":"All authors provided consent for publication of this manuscript.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}}]}}