{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T03:47:24Z","timestamp":1790048844069,"version":"4.0.1"},"reference-count":113,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2023,9,6]],"date-time":"2023-09-06T00:00:00Z","timestamp":1693958400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Research Council FORMAS (Project 2019-00288), Stockholm, Sweden","award":["PID2019-108320RJ-I00"],"award-info":[{"award-number":["PID2019-108320RJ-I00"]}]},{"name":"Research Council FORMAS (Project 2019-00288), Stockholm, Sweden","award":["IJCI-2015-24380"],"award-info":[{"award-number":["IJCI-2015-24380"]}]},{"name":"Research Council FORMAS (Project 2019-00288), Stockholm, Sweden","award":["RYC2020-028615-I"],"award-info":[{"award-number":["RYC2020-028615-I"]}]},{"DOI":"10.13039\/501100004837","name":"Spanish Ministry of Science and Innovation","doi-asserted-by":"publisher","award":["PID2019-108320RJ-I00"],"award-info":[{"award-number":["PID2019-108320RJ-I00"]}],"id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004837","name":"Spanish Ministry of Science and Innovation","doi-asserted-by":"publisher","award":["IJCI-2015-24380"],"award-info":[{"award-number":["IJCI-2015-24380"]}],"id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004837","name":"Spanish Ministry of Science and Innovation","doi-asserted-by":"publisher","award":["RYC2020-028615-I"],"award-info":[{"award-number":["RYC2020-028615-I"]}],"id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"publisher"}]},{"name":"FEDER funds (EU), Madrid, Spain","award":["PID2019-108320RJ-I00"],"award-info":[{"award-number":["PID2019-108320RJ-I00"]}]},{"name":"FEDER funds (EU), Madrid, Spain","award":["IJCI-2015-24380"],"award-info":[{"award-number":["IJCI-2015-24380"]}]},{"name":"FEDER funds (EU), Madrid, Spain","award":["RYC2020-028615-I"],"award-info":[{"award-number":["RYC2020-028615-I"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>The main cation\/calcium channel of spermatozoa (CatSper), first identified in 2001, has been thoroughly studied to elucidate its composition and function, while its distribution among species and sperm sources is yet incomplete. CatSper is composed of several subunits that build a pore-forming calcium channel, mainly activated in vivo in ejaculated sperm cells by intracellular alkalinization and progesterone, as suggested by the in vitro examinations. The CatSper channel relevance is dual: to maintain sperm homeostasis (alongside the plethora of membrane channels present) as well as being involved in pre-fertilization events, such as sperm capacitation, hyperactivation of sperm motility and the acrosome reaction, with remarkable species differences. Interestingly, the observed variations in CatSper localization in the plasma membrane seem to depend on the source of the sperm cells explored (i.e., epididymal or ejaculated, immature or mature, processed or not), the method used for examination and, particularly, on the specificity of the antibodies employed. In addition, despite multiple findings showing the relevance of CatSper in fertilization, few studies have studied CatSper as a biomarker to fine-tune diagnosis of sub-fertility in livestock or even consider its potential to control fertilization in plague animals, a more ethically defensible strategy than implicating CatSper to pharmacologically modify male-related fertility control in humans, pets or wild animals. This review describes inter- and intra-species differences in the localization, structure and function of the CatSper channel, calling for caution when considering its potential manipulation for fertility control or improvement.<\/jats:p>","DOI":"10.3390\/ijms241813750","type":"journal-article","created":{"date-parts":[[2023,9,6]],"date-time":"2023-09-06T10:07:37Z","timestamp":1693994857000},"page":"13750","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":13,"title":["The Cation\/Calcium Channel of Sperm (CatSper): A Common Role Played Despite Inter-Species Variation?"],"prefix":"10.3390","volume":"24","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0256-1958","authenticated-orcid":false,"given":"Alejandro","family":"Vicente-Carrillo","sequence":"first","affiliation":[{"name":"Department of Animal Production, Veterinary Faculty, Complutense University of Madrid, 28040 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0120-354X","authenticated-orcid":false,"given":"Manuel","family":"\u00c1lvarez-Rodr\u00edguez","sequence":"additional","affiliation":[{"name":"Department Animal Reproduction, Instituto Nacional de Investigaci\u00f3n y Tecnolog\u00eda Agraria y Alimentaria-Consejo Superior de Investigaciones Cient\u00edficas (INIA-CSIC), 28040 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5194-2124","authenticated-orcid":false,"given":"Heriberto","family":"Rodriguez-Martinez","sequence":"additional","affiliation":[{"name":"Department of Biomedical and Clinical Sciences (BKV), Link\u00f6ping University, 58225 Link\u00f6ping, Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,9,6]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Mann, T., and Lutwak-Mann, C. (1981). Male Reproductive Function and Semen, Springer. [1st ed.].","DOI":"10.1007\/978-1-4471-1300-3"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1071\/RD08183","article-title":"CatSper-null mutant spermatozoa are unable to ascend beyond the oviductal reservoir","volume":"21","author":"Ho","year":"2009","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_3","unstructured":"Vicente Carrillo, A. (2016). Sperm Membrane Channels, Receptors and Kinematics: Using Boar Spermatozoa for Drug Toxicity Screening. [PhD Thesis, Link\u00f6ping University]."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"473","DOI":"10.1387\/ijdb.082583bg","article-title":"Sperm head membrane reorganisation during capacitation","volume":"52","author":"Gadella","year":"2008","journal-title":"Int. J. Dev. Biol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"790","DOI":"10.1002\/j.1939-4640.2003.tb03123.x","article-title":"Tales from the tail: What do we really know about sperm motility?","volume":"24","author":"Turner","year":"2003","journal-title":"J. Androl."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"5623","DOI":"10.2741\/3105","article-title":"Ca2+ signalling in the control of motility and guidance in mammalian sperm","volume":"13","author":"Publicover","year":"2008","journal-title":"Front. Biosci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.theriogenology.2015.07.018","article-title":"An update on post-ejaculatory remodeling of the sperm surface before mammalian fertilization","volume":"85","author":"Gadella","year":"2016","journal-title":"Theriogenology"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1016\/j.ceca.2006.04.028","article-title":"T-type Ca2+ channels in sperm function","volume":"40","author":"Darszon","year":"2006","journal-title":"Cell Calcium."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1038\/nature09767","article-title":"Progesterone activates the principal Ca2+ channel of human sperm","volume":"471","author":"Lishko","year":"2011","journal-title":"Nature"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1111\/j.1365-2605.1990.tb00982.x","article-title":"Acidification of the epididymal fluid in the boar","volume":"13","author":"Ekstedt","year":"1990","journal-title":"Int. J. Androl."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"106829","DOI":"10.1016\/j.anireprosci.2021.106829","article-title":"How does the boar epididymis regulate the emission of fertile spermatozoa?","volume":"246","author":"Roca","year":"2022","journal-title":"Anim. Reprod. Sci."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Rodriguez-Martinez, H., Martinez, E.A., Calvete, J.J., Pe\u00f1a-Vega, F.J., and Roca, J. (2021). Seminal Plasma: Relevant for fertility?. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22094368"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"S138","DOI":"10.1016\/j.theriogenology.2007.03.018","article-title":"Role of the oviduct in sperm capacitation","volume":"68","year":"2007","journal-title":"Theriogenology"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1017\/S0967199406003777","article-title":"In vitro capacitation of bull spermatozoa by oviductal fluid and its components","volume":"3","author":"Bergqvist","year":"2006","journal-title":"Zygote"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1387\/ijdb.072554bn","article-title":"Ion channels that control fertility in mammalian spermatozoa","volume":"52","author":"Navarro","year":"2008","journal-title":"Int. J. Dev. Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"603","DOI":"10.1038\/35098027","article-title":"A sperm ion channel required for sperm motility and male fertility","volume":"413","author":"Ren","year":"2001","journal-title":"Nature"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1152\/physiol.00049.2009","article-title":"Calcium signalling through CatSper channels in mammalian fertilization","volume":"25","author":"Ren","year":"2010","journal-title":"Physiology"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"12527","DOI":"10.1073\/pnas.221454998","article-title":"A voltage-gated ion channel expressed specifically in spermatozoa","volume":"98","author":"Quill","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1480","DOI":"10.1016\/j.cell.2019.03.047","article-title":"Dual sensing of physiologic pH and calcium by EFCAB9 regulates sperm motility","volume":"177","author":"Hwang","year":"2019","journal-title":"Cell"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1186\/1477-7827-1-53","article-title":"Identification of human and mouse CATSPER3 and CATSPER4 genes: Characterisation of a common interaction domain and evidence for expression in testis","volume":"1","author":"Lobley","year":"2003","journal-title":"Reprod. Biol. Endocrinol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"18945","DOI":"10.1074\/jbc.M701083200","article-title":"CatSper\u03b2, a novel transmembrane protein in the CatSper channel complex","volume":"282","author":"Liu","year":"2007","journal-title":"J. Biol. Chem."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1095\/biolreprod.109.077107","article-title":"A novel, single, transmembrane protein CATSPERG is associated with CATSPER1 channel protein","volume":"81","author":"Wang","year":"2009","journal-title":"Biol. Reprod."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1038\/ncomms1153","article-title":"A novel gene required for male fertility and functional CATSPER channel formation in spermatozoa","volume":"2","author":"Chung","year":"2011","journal-title":"Nat. Commun."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"e23082","DOI":"10.7554\/eLife.23082","article-title":"CatSper\u03b6 regulates the structural continuity of sperm Ca2+ signalling domains and is required for normal fertility","volume":"6","author":"Chung","year":"2017","journal-title":"eLife"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"382","DOI":"10.1038\/nature09769","article-title":"The CatSper channel mediates progesterone-induced Ca2+ influx in human sperm","volume":"471","author":"Goodwin","year":"2011","journal-title":"Nature"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1152\/physiol.00028.2022","article-title":"CatSper Calcium Channels: 20 Years on","volume":"38","author":"Hwang","year":"2023","journal-title":"Physiology"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1095\/biolreprod.113.111708","article-title":"CatSper and the relationship of hyperactivated motility to intracellular calcium and pH kinetics in equine sperm","volume":"89","author":"Loux","year":"2013","journal-title":"Biol. Reprod."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"418","DOI":"10.1093\/humrep\/det454","article-title":"The CatSper calcium channel in human sperm: Relation with motility and involvement in progesterone-induced acrosome reaction","volume":"29","author":"Tamburrino","year":"2014","journal-title":"Hum. Reprod."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1532","DOI":"10.1093\/humrep\/dev103","article-title":"Quantification of CatSper1 expression in human spermatozoa and relation to functional parameters","volume":"30","author":"Tamburrino","year":"2015","journal-title":"Hum. Reprod."},{"key":"ref_30","first-page":"81","article-title":"Membrane receptor mapping in ejaculated ram spermatozoa","volume":"2105","author":"Casao","year":"2015","journal-title":"Reprod. Domest. Anim."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/j.repbio.2017.01.001","article-title":"The CatSper channel modulates boar sperm motility during capacitation","volume":"17","year":"2017","journal-title":"Reprod. Biol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1093\/biolre\/iox082","article-title":"Genomic identification, expression profiling, and functional characterization of CatSper channels in the bovine","volume":"97","author":"Johnson","year":"2017","journal-title":"Biol. Reprod."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"379","DOI":"10.15252\/embj.201489376","article-title":"The CatSper channel controls chemosensation in sea urchin sperm","volume":"34","author":"Seifert","year":"2015","journal-title":"EMBO J."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"102466","DOI":"10.1016\/j.ceca.2021.102466","article-title":"CatSper channels in sea urchin sperm","volume":"99","author":"Orta","year":"2021","journal-title":"Cell Calcium."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1038\/s41586-021-03742-6","article-title":"Structure of a mammalian sperm cation channel complex","volume":"595","author":"Lin","year":"2021","journal-title":"Nature"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"3439","DOI":"10.1038\/s41467-022-31050-8","article-title":"3D structure and in situ arrangements of CatSper channel in the sperm flagellum","volume":"13","author":"Zhao","year":"2022","journal-title":"Nat. Commun."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"808","DOI":"10.1016\/j.cell.2014.02.056","article-title":"Structurally distinct Ca(2+) signaling domains of sperm flagella orchestrate tyrosine phosphorylation and motility","volume":"157","author":"Chung","year":"2014","journal-title":"Cell"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"14869","DOI":"10.1073\/pnas.2136654100","article-title":"Hyperactivated sperm motility driven by CatSper2 is required for fertilization","volume":"100","author":"Quill","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1095\/biolreprod.107.060186","article-title":"CatSper3 and CatSper4 are essential for sperm hyperactivated motility and male fertility in the mouse","volume":"77","author":"Jin","year":"2007","journal-title":"Biol. Reprod."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1235","DOI":"10.1095\/biolreprod.105.045468","article-title":"CatSper3 and CatSper4 encode two cation channel-like proteins exclusively expressed in the testis","volume":"73","author":"Jin","year":"2005","journal-title":"Biol. Reprod."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1219","DOI":"10.1073\/pnas.0610286104","article-title":"All four CatSper ion channel proteins are required for male fertility and sperm cell hyperactivated motility","volume":"104","author":"Qi","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1007\/s10815-013-9972-6","article-title":"Levels of Tektin 2 and CatSper 2 in normozoospermic and oligoasthenozoospermic men and its association with motility, fertilization rate, embryo quality and pregnancy rate","volume":"30","author":"Bhilawadikar","year":"2013","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"6823","DOI":"10.1073\/pnas.1216588110","article-title":"Disruption of the principal, progesterone-activated sperm Ca2+ channel in a CatSper2-deficient infertile patient","volume":"110","author":"Smith","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"gaad015","DOI":"10.1093\/molehr\/gaad015","article-title":"SKF96365 modulates activity of CatSper channels in human sperm","volume":"29","author":"Brown","year":"2023","journal-title":"Mol. Hum. Reprod."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"13181","DOI":"10.1074\/jbc.RA120.013218","article-title":"The Ca2+ channel CatSper is not activated by cAMP\/PKA signaling but directly affected by chemicals used to probe the action of cAMP and PKA","volume":"295","author":"Wang","year":"2020","journal-title":"J. Biol. Chem."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1071\/RD06104","article-title":"State of the art in farm animal sperm evaluation","volume":"19","year":"2007","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"19546","DOI":"10.1038\/s41598-019-55834-z","article-title":"Release of Porcine Sperm from Oviduct Cells Is Stimulated by Progesterone and Requires CatSper","volume":"9","author":"Machado","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Mirihagalle, S., Hughes, J.R., and Miller, D.J. (2022). Progesterone-Induced Sperm Release from the Oviduct Sperm Reservoir. Cells, 11.","DOI":"10.3390\/cells11101622"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1111\/andr.13319","article-title":"Cortisol modulates Ca2+ signalling and acrosome reaction in human sperm","volume":"11","author":"Arias","year":"2023","journal-title":"Andrology"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Forero-Forero, A., L\u00f3pez-Ram\u00edrez, S., Felix, R., Hern\u00e1ndez-S\u00e1nchez, J., Tesoro-Cruz, E., Orozco-Su\u00e1rez, S., Murbarti\u00e1n, J., Soria-Castro, E., Olivares, A., and Bekker-M\u00e9ndez, C. (2022). Down Regulation of CatSper1 Expression by Calmodulin Inhibitor (Calmidazolium): Possible Implications for Fertility. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23158070"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"729","DOI":"10.1111\/andr.12054","article-title":"Intracellular calcium movements of boar spermatozoa during \u2018in vitro\u2019 capacitation and subsequent acrosome exocytosis follow a multiple storage place, extracellular calcium-dependent model","volume":"3","author":"Yeste","year":"2015","journal-title":"Andrology"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1530\/rep.1.00374","article-title":"Intracellular calcium and protein tyrosine phosphorylation during the release of bovine sperm adhering to the fallopian tube epithelium in vitro","volume":"129","author":"Gualtieri","year":"2005","journal-title":"Reproduction"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"5276","DOI":"10.1002\/jcp.27337","article-title":"Tyrosine phosphorylation signaling regulates Ca2+ entry by affecting intracellular pH during human sperm capacitation","volume":"234","author":"Bruckman","year":"2019","journal-title":"J. Cell Physiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"e13616","DOI":"10.1016\/j.heliyon.2023.e13616","article-title":"Hexane fraction of Prunus japonicahumbb. Seed extract enhances boar sperm motility via CatSper ion channel","volume":"9","author":"Cooray","year":"2023","journal-title":"Heliyon"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1007\/s13205-022-03452-4","article-title":"RNA-sequencing attest increased sperm motility in bovine spermatozoa treated with ethanolic extract of Putranjiva roxburghii","volume":"13","author":"Italiya","year":"2023","journal-title":"3 Biotech."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Ba\u0148as, \u0160., Benko, F., \u010eura\u010dka, M., Luk\u00e1\u010d, N., and Tvrd\u00e1, E. (2023). Epicatechin Prevents Cryocapacitation of Bovine Spermatozoa through Antioxidant Activity and Stabilization of Transmembrane Ion Channels. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24032510"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"e14512","DOI":"10.1111\/and.14512","article-title":"The effect of Fumaria parviflora on the expression of sexual hormones along with their receptors in testicles of adult rats induced by varicocele","volume":"54","author":"Dolatkhah","year":"2022","journal-title":"Andrologia"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"103465","DOI":"10.1016\/j.jtherbio.2023.103465","article-title":"Heat stress disrupts spermatogenesis via modulation of sperm-specific calcium channels in rats","volume":"112","author":"Ray","year":"2023","journal-title":"J. Therm. Biol."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Huang, X., Miyata, H., Wang, H., Mori, G., Iida-Norita, R., Ikawa, M., Percudani, R., and Chung, J.J. (2023). A CUG-initiated CATSPER\u03b8 functions in the CatSper channel assembly and serves as a checkpoint for flagellar trafficking. bioRxiv.","DOI":"10.1101\/2023.03.17.532952"},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Lacalle, E., Consuegra, C., Mart\u00ednez, C.A., Hidalgo, M., Dorado, J., Mart\u00ednez-Pastor, F., \u00c1lvarez-Rodr\u00edguez, M., and Rodr\u00edguez-Mart\u00ednez, H. (2022). Bicarbonate-Triggered In Vitro Capacitation of Boar Spermatozoa Conveys an Increased Relative Abundance of the Canonical Transient Receptor Potential Cation (TRPC) Channels 3, 4, 6 and 7 and of CatSper-\u03b3 Subunit mRNA Transcripts. Animals, 12.","DOI":"10.3390\/ani12081012"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1020628","DOI":"10.3389\/fmicb.2022.1020628","article-title":"Taxifolin increased semen quality of Duroc boars by improving gut microbes and blood metabolites","volume":"13","author":"Zhou","year":"2022","journal-title":"Front. Microbiol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/S0014-5793(03)00305-3","article-title":"Transient receptor potential (TRPC) channels in human sperm: Expression, cellular localization and involvement in the regulation of flagellar motility","volume":"541","author":"Castellano","year":"2003","journal-title":"FEBS Lett."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1111\/j.1748-1716.2011.02311.x","article-title":"Integration of transient receptor potential canonical channels with lipids","volume":"204","author":"Beech","year":"2012","journal-title":"Acta. Physiol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1305","DOI":"10.1152\/physrev.00028.2010","article-title":"Calcium channels in the development, maturation, and function of spermatozoa","volume":"91","author":"Darszon","year":"2011","journal-title":"Physiol. Rev."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1002\/j.1939-4640.1992.tb01631.x","article-title":"Movement characteristics of boar sperm obtained from the oviduct or hyperactivated in vitro","volume":"13","author":"Suarez","year":"1992","journal-title":"J. Androl."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1016\/S0378-4320(03)00134-9","article-title":"Sperm capacitation in the porcine oviduct","volume":"80","author":"Tienthai","year":"2004","journal-title":"Anim. Reprod. Sci."},{"key":"ref_67","unstructured":"Rodr\u00edguez-Mart\u00ednez, H. (1991). Aspects of the electrolytic composition of boar epididymal fluid with reference to sperm maturation and storage. Reprod. Domest. Anim., 13\u201327."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1590","DOI":"10.1095\/biolreprod.102.011320","article-title":"Characterization of the intracellular calcium store at the base of the sperm flagellum that regulates hyperactivated motility","volume":"68","author":"Ho","year":"2003","journal-title":"Biol. Reprod."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"652","DOI":"10.1017\/S096719941400029X","article-title":"ERK1\/2 mediates sperm acrosome reaction through elevation of intracellular calcium concentration","volume":"23","author":"Jaldety","year":"2015","journal-title":"Zygote"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1071\/BI9510581","article-title":"Observations on the penetration of the sperm in the mammalian egg","volume":"4","author":"Austin","year":"1951","journal-title":"Aust. J. Sci. Res. B"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"697","DOI":"10.1038\/168697b0","article-title":"Fertilizing capacity of spermatozoa deposited into the fallopian tubes","volume":"168","author":"Chang","year":"1951","journal-title":"Nature"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/S0304-4157(00)00018-6","article-title":"Dynamics of the mammalian sperm plasma membrane in the process of fertilization","volume":"1469","author":"Flesch","year":"2000","journal-title":"Biochim. Biophys. Acta."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"2407","DOI":"10.1242\/dev.127.11.2407","article-title":"The capacitating agent bicarbonate induces protein kinase A-dependent changes in phospholipid transbilayer behavior in the sperm plasma membrane","volume":"127","author":"Gadella","year":"2000","journal-title":"Development"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1002\/(SICI)1098-2795(200002)55:2<220::AID-MRD12>3.0.CO;2-I","article-title":"cAMP-dependent protein kinase control of plasma membrane lipid architecture in boar sperm","volume":"55","author":"Harrison","year":"2000","journal-title":"Mol. Reprod. Dev."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"816","DOI":"10.1038\/aja.2012.81","article-title":"Mechanism of sperm capacitation and the acrosome reaction: Role of protein kinases","volume":"14","author":"Ickowicz","year":"2012","journal-title":"Asian. J. Androl."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1530\/jrf.0.0230193","article-title":"The movement of golden hamster spermatozoa before and after capacitation","volume":"23","author":"Yanagimachi","year":"1970","journal-title":"J. Reprod. Fertil."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"823","DOI":"10.1016\/j.tibs.2019.08.001","article-title":"Tiny Dancer: EFCAB9 Triggers Sperm Hyperactivation via CatSper","volume":"44","author":"Bainbridge","year":"2019","journal-title":"Trends Biochem. Sci."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"1178","DOI":"10.1038\/ejhg.2010.108","article-title":"Genetic male infertility and mutation of CATSPER ion channels","volume":"18","author":"Hildebrand","year":"2010","journal-title":"Eur. J. Hum. Genet."},{"key":"ref_79","first-page":"2737","article-title":"Specific loss of CatSper function is sufficient to compromise fertilizing capacity of human spermatozoa","volume":"30","author":"Williams","year":"2015","journal-title":"Hum. Reprod."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"dev199988","DOI":"10.1242\/dev.199988","article-title":"C2CD6 regulates targeting and organization of the CatSper calcium channel complex in sperm flagella","volume":"149","author":"Yang","year":"2022","journal-title":"Development"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1958","DOI":"10.1016\/j.theriogenology.2016.06.016","article-title":"Melatonin receptors MT1 and MT2 are expressed in spermatozoa from several seasonal and nonseasonal breeder species","volume":"86","author":"Roca","year":"2016","journal-title":"Theriogenology"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1095\/biolreprod21.1.173","article-title":"Movements of sodium and potassium into epididymal boar spermatozoa","volume":"21","author":"Zimmerman","year":"1979","journal-title":"Biol. Reprod."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1071\/RD05120","article-title":"Moving to the beat: A review of mammalian sperm motility regulation","volume":"18","author":"Turner","year":"2006","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1016\/j.ydbio.2006.11.007","article-title":"Contributions of extracellular and intracellular Ca2+ to regulation of sperm motility: Release of intracellular stores can hyperactivate CatSper1 and CatSper2 null sperm","volume":"303","author":"Marquez","year":"2007","journal-title":"Dev. Biol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1017\/S1464793103006407","article-title":"The sperm, a neuron with a tail: \u201cneurona\u201d receptors in mammalian sperm","volume":"79","author":"Meizel","year":"2004","journal-title":"Biol. Rev. Camb. Philos. Soc."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1111\/rda.12728","article-title":"Membrane Stress During Thawing Elicits Redistribution of Aquaporin 7 But Not of Aquaporin 9 in Boar Spermatozoa","volume":"51","author":"Ekwall","year":"2016","journal-title":"Reprod. Domest. Anim."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Parra, A., Padilla, L., Lucas, X., Rodriguez-Martinez, H., Barranco, I., and Roca, J. (2023). Seminal extracellular vesicles and their involvement in male (in)fertility: A systematic review. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24054818"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.1071\/RD9961165","article-title":"Viability and membrane integrity of spermatozoa after dilution and flow cytometric sorting in the presence or absence of seminal plasma","volume":"8","author":"Maxwell","year":"1996","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_89","first-page":"1","article-title":"The physiological roles of the boar ejaculate","volume":"66","author":"Kvist","year":"2009","journal-title":"Soc. Reprod. Fertil. Suppl."},{"key":"ref_90","first-page":"221","article-title":"Characteristics of sperm motility in boar semen diluted in different extenders and stored for seven days at 18 degrees C","volume":"7","author":"Estienne","year":"2007","journal-title":"Reprod. Biol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1016\/j.theriogenology.2015.07.009","article-title":"Artificial insemination in pigs today","volume":"85","author":"Knox","year":"2016","journal-title":"Theriogenology"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"100608","DOI":"10.1016\/j.mtbio.2023.100608","article-title":"Exosomes: New regulators of reproductive development","volume":"19","author":"Chen","year":"2023","journal-title":"Mater Today Bio."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"106853","DOI":"10.1016\/j.anireprosci.2021.106853","article-title":"Extracellular vesicles in seminal fluid and its impact on male reproduction. An overview in pet and livestock species","volume":"246","author":"Roca","year":"2022","journal-title":"Anim. Reprod. Sci."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"107051","DOI":"10.1016\/j.anireprosci.2022.107051","article-title":"Extracellular vesicles in seminal plasma: A safe and relevant tool to improve fertility in livestock?","volume":"244","author":"Roca","year":"2022","journal-title":"Anim. Reprod. Sci."},{"key":"ref_95","doi-asserted-by":"crossref","unstructured":"Zhang, X., Song, D., Kang, H., Zhou, W., Chen, H., and Zeng, X. (2020). Seminal plasma exosomes evoke calcium signals via the CatSper channel to regulate human sperm function. bioRxiv.","DOI":"10.1101\/2020.05.21.094433"},{"key":"ref_96","doi-asserted-by":"crossref","unstructured":"Zhang, X., Liang, M., Song, D., Huang, R., Chen, C., Chen, H., Wang, Q., Sun, X., Song, J., and Zhang, J. (2023). Extracellular vesicles of seminal plasma and their derived peptides improve human sperm function via the CatSper-mediated calcium signals. Lancet, prepint.","DOI":"10.2139\/ssrn.4456595"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"100514","DOI":"10.1016\/j.mcpro.2023.100514","article-title":"The proteome of large or small extracellular vesicles in pig seminal plasma differs, defining sources and biological functions","volume":"22","author":"Barranco","year":"2022","journal-title":"Mol. Cell. Proteomics."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"1102049","DOI":"10.3389\/fvets.2023.1102049","article-title":"Extracellular vesicles would be involved in the release and delivery of seminal TGF-\u03b2 isoforms in pigs","volume":"10","author":"Padilla","year":"2023","journal-title":"Front. Vet. Sci."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1186\/1477-7827-2-75","article-title":"Role of tyrosine phosphorylation in sperm capacitation\/acrosome reaction","volume":"2","author":"Naz","year":"2004","journal-title":"Reprod. Biol. Endocrinol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1186\/s12958-017-0281-2","article-title":"The CatSper channel and its roles in male fertility: A systematic review","volume":"15","author":"Sun","year":"2017","journal-title":"Reprod. Biol. Endocrinol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1051\/rnd:2000120","article-title":"The physiological role of beta-endorphin in porcine ovarian follicles","volume":"40","author":"Kaminski","year":"2000","journal-title":"Reprod. Nutr. Dev."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"846","DOI":"10.2119\/molmed.2010.00268","article-title":"Regulation of male fertility by the opioid system","volume":"17","author":"Casis","year":"2011","journal-title":"Mol. Med."},{"key":"ref_103","first-page":"131","article-title":"Metabolic control through L calcium channel, PKC and opioid receptors modulation by an association of naloxone and calcium salts. Curr Drug Targets Immune","volume":"1","author":"Minoia","year":"2001","journal-title":"Endocr. Metabol. Disord."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1146\/annurev.pharmtox.40.1.389","article-title":"Molecular mechanisms and regulation of opioid receptor signaling","volume":"40","author":"Law","year":"2000","journal-title":"Annu. Rev. Pharmacol. Toxicol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1659","DOI":"10.1016\/S0024-3205(97)00108-2","article-title":"Effects of naloxone on amphetamine induced striatal dopamine release in vivo: A microdialysis study","volume":"60","author":"Feigenbaum","year":"1997","journal-title":"Life. Sci."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"724","DOI":"10.1002\/mrd.22675","article-title":"The mu (\u03bc) and delta (\u03b4) opioid receptors modulate boar sperm motility","volume":"83","year":"2016","journal-title":"Mol. Reprod. Dev."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1373\/clinchem.2018.295089","article-title":"Male Contraceptive Development: Update on Novel Hormonal and Nonhormonal Methods","volume":"65","author":"Long","year":"2019","journal-title":"Clin. Chem."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Mariani, N.A.P., Silva, J.V., Fardilha, M., and Silva, E.J.R. (2023). Advances in non-hormonal male contraception targeting sperm motility. Hum. Reprod. Update, dmad008.","DOI":"10.1093\/humupd\/dmad008"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"1010306","DOI":"10.3389\/fcell.2023.1010306","article-title":"High-throughput screening method for discovering CatSper inhibitors using membrane depolarization caused by external calcium chelation and fluorescent cell barcoding","volume":"11","author":"Luque","year":"2023","journal-title":"Front. Cell. Dev. Biol."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"758","DOI":"10.1093\/humupd\/dmz032","article-title":"Human sperm ion channel (dys)function: Implications for fertilization","volume":"25","author":"Brown","year":"2019","journal-title":"Hum. Reprod. Update"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"3144","DOI":"10.1111\/bph.14355","article-title":"A novel cross-species inhibitor to study the function of CatSper Ca2+ channels in sperm","volume":"175","author":"Rennhack","year":"2018","journal-title":"Br. J. Pharmacol."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"e202000499","DOI":"10.1002\/cmdc.202000499","article-title":"Discovery and Characterization of Multiple Classes of Human CatSper Blockers","volume":"17","author":"Carlson","year":"2022","journal-title":"Chem. Med. Chem."},{"key":"ref_113","doi-asserted-by":"crossref","unstructured":"Cai, X., and Clapham, D.E. (2008). Evolutionary genomics reveals lineage-specific gene loss and rapid evolution of a sperm-specific ion channel complex: CatSpers and CatSperbeta. PLoS ONE, 3.","DOI":"10.1371\/journal.pone.0003569"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/24\/18\/13750\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T20:46:09Z","timestamp":1760129169000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/24\/18\/13750"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,9,6]]},"references-count":113,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2023,9]]}},"alternative-id":["ijms241813750"],"URL":"https:\/\/doi.org\/10.3390\/ijms241813750","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,9,6]]}}}