Animal Reproduction (AR)
https://animal-reproduction.org/article/doi/10.1590/1984-3143-AR2026-0051
Animal Reproduction (AR)
Thematic Section: 42nd Annual Scientific Meeting of the Association of Embryo Technology in Europe (AETE)

Biomarkers of sperm fertility in pigs: a relevant tool for the discrimination of sub‑fertile males?

Manuel Álvarez-Rodríguez; Adrián Martín-San Juan; Helena Nieto-Cristóbal; Alejandro Vicente-Carrillo; Eduardo de Mercado

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Abstract

In internally fertilizing animals, such as the pig, males transfer not only spermatozoa but also a complex mixture collectively termed seminal plasma (SP). SP contains diverse bioactive components, including proteins, peptides, cytokines, and various RNA species. Growing evidence indicates that SP contributes not only to the sperm quality but also modulates the female’s immune responses. Major progress in biological knowledge and in the discovery of sperm quality biomarkers has been driven by advances in biotechnology and by the increasing affordability of omics technologies. With modern proteomic, (epi)genomic, transcriptomic, metabolomic, and functional analyses applied directly to semen, it is now possible to evaluate not only the functional state of the testes, epididymis, and accessory glands, but also to predict fertility through emerging biomarkers. Beyond diagnostics and fertility forecasting, new opportunities arise from the sperm quality biomarker definition using semen additives—such as specific molecules (proteins, peptides, enzymes) or extracellular vesicles (EVs). These EVs contain bioactive cargo protected within nanovesicles that can be isolated, stored, or even produced de novo in vitro. Boar spermatozoa are notoriously difficult to cryopreserve because they lose cholesterol during handling. EVs could fuse with sperm cells, deliver cholesterol‑rich lipids to the sperm membrane, and transfer specific proteins and nucleic acids. This raises the possibility that harvesting EVs, especially from highly fertile males, could be used in the future to improve sperm storage and even cryopreservation. Taken together, this review aims to list sperm fertility biomarkers that could be useful for the AI companies to discriminate sub-fertile males at an early stage.

Keywords

extracellular vesicles, receptors, sperm physiology, fertility

References

Alegre E, Biehl M, Petkov N, Sánchez L. Automatic classification of the acrosome status of boar spermatozoa using digital image processing and LVQ. Comput Biol Med. 2008;38(4):461-8. https://doi.org/10.1016/j.compbiomed.2008.01.005. PMid:18339365.

Alminana C, Caballero I, Heath PR, Maleki-Dizaji S, Parrilla I, Cuello C, Gil MA, Vazquez JL, Vazquez JM, Roca J, Martinez EA, Holt WV, Fazeli A. The battle of the sexes starts in the oviduct: modulation of oviductal transcriptome by X and Y-bearing spermatozoa. BMC Genomics. 2014;15(1):293. https://doi.org/10.1186/1471-2164-15-293. PMid:24886317.

Alvarez-Rodriguez M, Atikuzzaman M, Venhoranta H, Wright D, Rodriguez-Martinez H. Expression of immune regulatory genes in the porcine internal genital tract is differentially triggered by spermatozoa and seminal plasma. Int J Mol Sci. 2019;20(3):1-20. https://doi.org/10.3390/ijms20030513. PMid:30691059.

Alvarez-Rodriguez M, Martinez C, Wright D, Barranco I, Roca J, Rodriguez-Martinez H. The transcriptome of pig spermatozoa, and its role in fertility. Int J Mol Sci. 2020a;21(5):1572. https://doi.org/10.3390/ijms21051572. PMid:32106598.

Alvarez-Rodriguez M, Martinez CA, Roca J, Rodriguez-Martinez H. mRNA expression of oxidative-reductive proteins in boars with documented different fertility can identify relevant prognostic biomarkers. Res Vet Sci. 2021;141:195-202. https://doi.org/10.1016/j.rvsc.2021.10.022. PMid:34763256.

Álvarez-Rodríguez M, Martinez CA, Wright D, Rodríguez-Martinez H. The role of semen and seminal plasma in inducing large-scale genomic changes in the female porcine peri-ovulatory tract. Sci Rep. 2020;10(1):5061. https://doi.org/10.1038/s41598-020-60810-z. PMid:32193402.

Alvarez-Rodriguez M, Martinez CA, Wright D, Rodriguez-Martinez H. Does the act of copulation per se, without considering seminal deposition, change the expression of genes in the porcine female genital tract? Int J Mol Sci. 2020b;21(15):5477. https://doi.org/10.3390/ijms21155477. PMid:32751869.

Amann RP, Katz DF. Andrology lab corner: reflections on CASA after 25 years. J Androl. 2004;25(3):317-25. https://doi.org/10.1002/j.1939-4640.2004.tb02793.x. PMid:15064305.

Atikuzzaman M, Alvarez-Rodriguez M, Vicente-Carrillo A, Johnsson M, Wright D, Rodriguez-Martinez H. Conserved gene expression in sperm reservoirs between birds and mammals in response to mating. BMC Genomics. 2017;18(1):98. https://doi.org/10.1186/s12864-017-3488-x. PMid:28100167.

Atikuzzaman M, Hoglund A, Johnsson M, Alvarez-Rodriguez M, Mehta R, Wright D, Rodriguez-Martinez H. Spermatozoa rather than seminal fluid drives immune-modulation in the sperm storage tubuli of Red Jungle Fowl hens. Reprod Domest Anim. 2015

Ausejo-Marcos R, Miguel-Jiménez S, Tejedor MT, Gómez-Giménez B, Soriano-Úbeda C, Martinez-Pastor F, Mendoza N, Vicente-Carrillo A, Hurtado WF, Holguín CÁ, Moreno B, Falceto MV. Acrosomal status and PARP-1 nuclear markers could improve discrimination of potential fertility in good-quality boar semen doses. Reprod Domest Anim. 2025;60(11):e70145. https://doi.org/10.1111/rda.70145. PMid:41222051.

Ax RL, Dally M, Didion BA, Lenz RW, Love CC, Varner DD, Hafez B, Bellin ME. Semen evaluation. In: Hafez B, Hafez ESE, editors. Reproduction in farm animals. Hoboken: Wiley; 2000. p. 363–75. https://doi.org/10.1002/9781119265306.ch25.

Bae J-W, Hwang J-M, Kwon W-S. Prediction of male fertility using Ras-related proteins. J Anim Sci Technol. 2022;64(6):1024-34. https://doi.org/10.5187/jast.2022.e83. PMid:36812003.

Bae J-W, Hwang J-M, Lee W-J, Kim D-H, Yi JK, Ha JJ, Oh DY, Kwon W-S. Application of sperm motion kinematics and motility-related proteins for prediction of male fertility. Theriogenology. 2024;218:223-30. https://doi.org/10.1016/j.theriogenology.2024.02.007. PMid:38359560.

Bae J-W, Kim S-H, Kim D-H, Ha JJ, Yi JK, Hwang S, Ryu B-Y, Pang M-G, Kwon W-S. Ras-related proteins (Rab) are key proteins related to male fertility following a unique activation mechanism. Reprod Biol. 2019;19(4):356-62. https://doi.org/10.1016/j.repbio.2019.10.001. PMid:31606348.

Baker RD, Degen AA. Transport of live and dead boar spermatozoa within the reproductive tract of gilts. J Reprod Fertil. 1972;28(3):369-77. https://doi.org/10.1530/jrf.0.0280369. PMid:4111336.

Barranco I, Fernandez-Fuertes B, Padilla L, Delgado-Bermúdez A, Tvarijonaviciute A, Yeste M. Seminal plasma anti-müllerian hormone: a potential ai-boar fertility biomarker? Biology (Basel). 2020;9(4):78. https://doi.org/10.3390/biology9040078. PMid:32290279.

Barranco I, Martínez-Díaz P, Parra A, Martínez-Alborcia MJ, Lucas X, Rodríguez-Martínez H, Roca J. Proteomic profiling of porcine seminal extracellular vesicles reveals potential in vivo fertility biomarkers. Andrology. 2026;14(2):555-70. https://doi.org/10.1111/andr.70089. PMid:40613678.

Barranco I, Tvarijonaviciute A, Perez-Patinõ C, Parrilla I, Ceron JJ, Martinez EA, Rodriguez-Martinez H, Roca J. High total antioxidant capacity of the porcine seminal plasma (SP-TAC) relates to sperm survival and fertility. Sci Rep. 2015;5(1):18538. https://doi.org/10.1038/srep18538. PMid:26688188.

Barranco S, Tvarijonaviciute A, Perez-Patiho C, Vicente-Carrillo A, Parrilla N, Ceron JJ, Martinez EA, Rodriguez-Martinez H, Roca J. Glutathione peroxidase 5 is expressed by the entire pig male genital tract and once in the seminal plasma contributes to sperm survival and in vivo fertility. PLoS One. 2016;11(9):e0162958. https://doi.org/10.1371/journal.pone.0162958. PMid:27627110.

Barraud-Lange V, Chalas Boissonnas C, Serres C, Auer J, Schmitt A, Lefèvre B, Wolf J-P, Ziyyat A. Membrane transfer from oocyte to sperm occurs in two CD9-independent ways that do not supply the fertilising ability of Cd9-deleted oocytes. Reproduction. 2012;144(1):53-66. https://doi.org/10.1530/REP-12-0040. PMid:22554680.

Barraud-Lange V, Naud-Barriant N, Bomsel M, Wolf J-P, Ziyyat A. Transfer of oocyte membrane fragments to fertilizing spermatozoa. FASEB J. 2007a;21(13):3446-9. https://doi.org/10.1096/fj.06-8035hyp. PMid:17575263.

Barraud-Lange V, Naud-Barriant N, Saffar L, Gattegno L, Ducot B, Drillet A-S, Bomsel M, Wolf J-P, Ziyyat A. Alpha6beta1 integrin expressed by sperm is determinant in mouse fertilization. BMC Dev Biol. 2007b;7(1):102. https://doi.org/10.1186/1471-213X-7-102. PMid:17850654.

Batista C, van Lier E, Petrocelli H. Dynamics of sperm DNA fragmentation in raw boar semen and fertility. Reprod Domest Anim. 2016;51(5):774-80. https://doi.org/10.1111/rda.12749. PMid:27546051.

Björndahl L, Brown JK. The sixth edition of the WHO Laboratory Manual for the Examination and Processing of Human Semen: ensuring quality and standardization in basic examination of human ejaculates. Fertility Sterility. 2022;117(2):246-51. https://doi.org/10.1016/j.fertnstert.2021.12.012.

Boe-Hansen GB, Christensen P, Vibjerg D, Nielsen MBF, Hedeboe AM. Sperm chromatin structure integrity in liquid stored boar semen and its relationships with field fertility. Theriogenology. 2008;69(6):728-36. https://doi.org/10.1016/j.theriogenology.2007.12.004. PMid:18242673.

Boe-Hansen GB, Satake N. An update on boar semen assessments by flow cytometry and CASA. Theriogenology. 2019;137:93-103. https://doi.org/10.1016/j.theriogenology.2019.05.043. PMid:31200933.

Braundmeier AG, Demers JM, Shanks RD, Miller DJ. The relationship of porcine sperm zona-binding ability to fertility. J Anim Sci. 2004;82(2):452-8. https://doi.org/10.2527/2004.822452x. PMid:14974543.

Brinke I, Große-Brinkhaus C, Roth K, Pröll-Cornelissen MJ, Klein S, Schellander K, Tholen E. Endocrine fertility parameters-genomic background and their genetic relationship to boar taint in german landrace and large white. Animals (Basel). 2021;11(1):231. https://doi.org/10.3390/ani11010231. PMid:33477702.

Broekhuijse MLWJ, Feitsma H, Gadella BM. Artificial insemination in pigs: predicting male fertility. Vet Q. 2012;32(3-4):151-7. https://doi.org/10.1080/01652176.2012.735126. PMid:23092203.

Broekhuijse MLWJ, Feitsma H, Gadella BM. Field data analysis of boar semen quality. Reprod Domest Anim. 2011;46(s2 Suppl 2):59-63. https://doi.org/10.1111/j.1439-0531.2011.01861.x. PMid:21884280.

Broekhuijse MLWJ, Šoštarić E, Feitsma H, Gadella BM. The value of microscopic semen motility assessment at collection for a commercial artificial insemination center, a retrospective study on factors explaining variation in pig fertility. Theriogenology. 2012a;77(7):1466-1479.e3. https://doi.org/10.1016/j.theriogenology.2011.11.016. PMid:22289218.

Broekhuijse MLWJ, Šoštarić E, Feitsma H, Gadella BM. Application of computer-assisted semen analysis to explain variations in pig fertility. J Anim Sci. 2012b;90(3):779-89. https://doi.org/10.2527/jas.2011-4311. PMid:22064743.

Bu Y, Wang P, Li S, Li L, Zhang S, Wei H. Semen protein CRISP3 promotes reproductive performance of boars through immunomodulation. Int J Mol Sci. 2024;25(4):2264. https://doi.org/10.3390/ijms25042264. PMid:38396941.

Caballero I, Vazquez JM, Mayor GM, Almiñana C, Calvete JJ, Sanz L, Roca J, Martinez EA. PSP-I/PSP-II spermadhesin exert a decapacitation effect on highly extended boar spermatozoa. Int J Androl. 2009;32(5):505-13. https://doi.org/10.1111/j.1365-2605.2008.00887.x. PMid:18399981.

Chen G, Wang Y, Kang J, Mei X, Zhang Z, Zhang Y, Lu J, Liu Y, Lan F, Huang W, Zhang D. CFAP74 variants could cause male infertility with the asthenoteratozoospermia phenotype. Ann Hum Genet. 2025a. https://doi.org/10.1111/ahg.70026. PMid:41117181.

Chen J, Huang R, Ma J, Su G, Huang M, Zhou W, Liu C, Liu Q, Li P, Zhao Q. Investigating the genetic imprint of long body length, high lean meat rate, high fertility and long gestation period in Danish Landrace pigs. BMC Genomics. 2025b;26(1):869. https://doi.org/10.1186/s12864-025-12092-w. PMid:41023780.

Chen J, Lin X, Bhattacharya S, Wiesehöfer C, Wennemuth G, Müller K, Montag D. Neuroplastin expression in male mice is essential for fertility, mating, and adult testosterone levels. Int J Mol Sci. 2023;25(1):177. https://doi.org/10.3390/ijms25010177. PMid:38203350.

Chen PR, Redel BK, Kerns KC, Spate LD, Prather RS. Challenges and considerations during in vitro production of porcine embryos. Cells. 2021;10(10):2770. https://doi.org/10.3390/cells10102770. PMid:34685749.

Chen W, Xie Y, Xu Z, Shang Y, Yang W, Wang P, Wu Z, Cai G, Hong L. Identification and functional analysis of miRNAs in extracellular vesicles of semen plasma from high- and low-fertility boars. Animals (Basel). 2024;15(1):40. https://doi.org/10.3390/ani15010040. PMid:39794983.

Cheng J, Hao X, Zhang W, Sun C, Yuan X, Yang Y, Zeng W, Zhu Z. Proteomic and metabolomic profiling reveals alterations in boar X and Y sperm. Animals (Basel). 2024;14(24):3672. https://doi.org/10.3390/ani14243672. PMid:39765576.

Choi H, Han C, Jin S, Kwon JT, Kim J, Jeong J, Kim J, Ham S, Jeon S, Yoo YJ, Cho C. Reduced fertility and altered epididymal and sperm integrity in mice lacking ADAM71. Biol Reprod. 2015;93(3):70. https://doi.org/10.1095/biolreprod.115.130252. PMid:26246218.

Collins ED, Flowers WL, Shanks RD, Miller DJ. Porcine sperm zona binding ability as an indicator of fertility. Anim Reprod Sci. 2008;104(1):69-82. https://doi.org/10.1016/j.anireprosci.2007.01.012. PMid:17303353.

Congras A, Yerle-Bouissou M, Pinton A, Vignoles F, Liaubet L, Ferchaud S, Acloque H. Sperm DNA methylation analysis in swine reveals conserved and species-specific methylation patterns and highlights an altered methylation at the GNAS locus in infertile boars. Biol Reprod. 2014;91(6):137. https://doi.org/10.1095/biolreprod.114.119610. PMid:25320151.

Daigneault BW, McNamara KA, Purdy PH, Krisher RL, Knox RV, Rodriguez-Zas SL, Miller DJ. Enhanced fertility prediction of cryopreserved boar spermatozoa using novel sperm function assessment. Andrology. 2015;3(3):558-68. https://doi.org/10.1111/andr.12035. PMid:25914302.

de Mercado E, Nieto-Cristóbal H, Martín-San Juan A, Martinez-Alborcia MJ, Álvarez-Rodríguez M. Could individual variability in resistance to cryopreservation (“freezability”) serve as a biomarker reflecting boar fertility? Animals (Basel). 2025;15(15):2180. https://doi.org/10.3390/ani15152180. PMid:40804970.

Desaulniers AT, Ross CE, White BR. Porcine leydig cells: central regulators of boar reproductive development and fertility. J Anim Sci. 2026;104:skag215. https://doi.org/10.1093/jas/skag215. PMid:42434794.

Didion BA, Braun GD, Duggan MV. Field fertility of frozen boar semen: A retrospective report comprising over 2600 AI services spanning a four year period. Anim Reprod Sci. 2013;137(3-4):189-96. https://doi.org/10.1016/j.anireprosci.2013.01.001. PMid:23348011.

Didion BA, Kasperson KM, Wixon RL, Evenson DP. Boar fertility and sperm chromatin structure status: a retrospective report. J Androl. 2009;30(6):655-60. https://doi.org/10.2164/jandrol.108.006254. PMid:19478334.

Druart X, Gatti J-L, Huet S, Dacheux J-L, Humblot P. Hypotonic resistance of boar spermatozoa: sperm subpopulations and relationship with epididymal maturation and fertility. Reproduction. 2009;137(2):205-13. https://doi.org/10.1530/REP-08-0225. PMid:18996973.

Edwards DR, Handsley MM, Pennington CJ. The ADAM metalloproteinases. Mol Aspects Med. 2009. https://doi.org/10.1016/j.mam.2008.08.001. PMid:18762209.

Fernández-López P, Garriga J, Casas I, Yeste M, Bartumeus F. Predicting fertility from sperm motility landscapes. Commun Biol. 2022;5(1):1027. https://doi.org/10.1038/s42003-022-03954-0. PMid:36171267.

Flowers WL. Selection for boar fertility and semen quality--the way ahead. Soc Reprod Fertil Suppl. 2009;66:67-78. PMid:19848267.

Foxcroft GR, Patterson J, Cameron A, Dyck MK. Application of advanced AI technologies to improve the competitiveness of the pork industry. IPVS Congr.2010;2010:25-29.

Gadea J. Sperm factors related to in vitro and in vivo porcine fertility. Theriogenology. 2005;63(2):431-44. https://doi.org/10.1016/j.theriogenology.2004.09.023. PMid:15626409.

Gadea J, Matás C, Lucas X. Prediction of porcine semen fertility by homologous in vitro penetration (hIVP) assay. Anim Reprod Sci. 1998;54(2):95-108. https://doi.org/10.1016/S0378-4320(98)00144-4. PMid:9877056.

Gadea J, Sellés E, Marco MA. The predictive value of porcine seminal parameters on fertility outcome under commercial conditions. Reprod Domest Anim. 2004;39(5):303-8. https://doi.org/10.1111/j.1439-0531.2004.00513.x. PMid:15367261.

Gao F, Wang P, Wang K, Fan Y, Chen Y, Chen Y, Ye C, Feng M, Li L, Zhang S, Wei H. Investigation into the relationship between sperm Cysteine-Rich Secretory Protein 2 (CRISP2) and sperm fertilizing ability and fertility of boars. Front Vet Sci. 2021;8:653413. https://doi.org/10.3389/fvets.2021.653413. PMid:33996980.

Garcia-Canovas M, Parrilla I, Cuello C, Gil MA, Martinez EA. Swine in vitro embryo production: Potential, challenges, and advances. Anim Reprod Sci. 2024;270:107600. https://doi.org/10.1016/j.anireprosci.2024.107600. PMid:39270509.

Gardela J, Ruiz-Conca M, Martinez CA, Wright D, López-Béjar M, Rodriguez-Martinez H, Alvarez-Rodriguez M. The expression of cold-inducible RNA-binding protein mRNA in sow genital tract is modulated by natural mating, but not by seminal plasma. Int J Mol Sci. 2020;21(15):5333. https://doi.org/10.3390/ijms21155333. PMid:32727091.

Gershoni M, Hauser R, Yogev L, Lehavi O, Azem F, Yavetz H, Pietrokovski S, Kleiman SE. A familial study of azoospermic men identifies three novel causative mutations in three new human azoospermia genes. Genet Med. 2017;19(9):998-1006. https://doi.org/10.1038/gim.2016.225. PMid:28206990.

Ghanbari H, Keshtgar S, Zare HR, Gharesi-Fard B. Inhibition of CatSper and Hv1 channels and NOX5 enzyme affect progesterone-induced increase of intracellular calcium concentration and ROS generation in human sperm. Iran J Med Sci. 2019;44(2):127-34. PMid:30936599.

Gòdia M, Reverter A, González-Prendes R, Ramayo-Caldas Y, Castelló A, Rodríguez-Gil J-E, Sánchez A, Clop A. A systems biology framework integrating GWAS and RNA-seq to shed light on the molecular basis of sperm quality in swine. Genet Sel Evol. 2020;52(1):72. https://doi.org/10.1186/s12711-020-00592-0. PMid:33292187.

Han M, Yao D, Song Y, Liu Y, Chen Z, Li J, Li F, Yang X, Dai L, Niu B. Identification of functional SNP associated with sperm quality in porcine ANXA5 that contributes to the growth of immature Sertoli cell. Front Vet Sci. 2025;12:1576566. https://doi.org/10.3389/fvets.2025.1576566. PMid:40438404.

Harrison RA. Sperm plasma membrane characteristics and boar semen fertility. J Reprod Fertil Suppl. 1997;52:195-211. PMid:9602729.

Henneberg S, Pieper L, Selige C, Jung M, Schulze M. Analysis of artificial insemination center management factors that contribute to sperm parameters and boar longevity with a major focus on PGF2α treatment. J Anim Sci. 2023;101:skad251. https://doi.org/10.1093/jas/skad251. PMid:37523262.

Hensel B, Henneberg S, Kleve-Feld M, Jung M, Schulze M. Selection and direct biomarkers of reproductive capacity of breeding boars. Anim Reprod Sci. 2024;269:107490. https://doi.org/10.1016/j.anireprosci.2024.107490. PMid:38735766.

Hensel B, Henneberg S, Marini S, Jung M, Schulze M. Artificial intelligence-based prediction of boar reproductive fitness and health: current status in research and practice. Anim Reprod Sci. 2026;289:108172. https://doi.org/10.1016/j.anireprosci.2026.108172. PMid:41861691.

Herrera J, Fierro R, Zayas H, Conejo J, Jiménez I, García A, Betancourt M. Acrosome reaction in fertile and subfertile boar sperm. Arch Androl. 2002;48(2):133-9. https://doi.org/10.1080/014850102317267445. PMid:11868626.

Holt C, Holt WV, Moore HD, Reed HC, Curnock RM. Objectively measured boar sperm motility parameters correlate with the outcomes of on-farm inseminations: results of two fertility trials. J Androl. 1997;18(3):312-23. https://doi.org/10.1002/j.1939-4640.1997.tb01925.x. PMid:9203061.

Huang J, Zuo Z, Zhao H, Wang C, Li S, Liu Z, Yang Y, Jiang S. Cluster analysis and potential influencing factors of boars with different fertility. Theriogenology. 2023;199:95-105. https://doi.org/10.1016/j.theriogenology.2022.12.039. PMid:36709653.

Imran M, Pettitt M, Buhr MM. Characterization of Na+⁄K+-ATPase associated with boar fertility. Theriogenology. 2025;243:117460. https://doi.org/10.1016/j.theriogenology.2025.117460. PMid:40373676.

Jang S-I, Jo J-H, Claudine U, Jung E-J, Lee W-J, Hwang J-M, Bae J-W, Kim D-H, Yi JK, Ha JJ, Oh DY, Kwon W-S. Correlation between Rab3A Expression and Sperm Kinematic Characteristics. Dev Reprod. 2024;28(1):13-9. https://doi.org/10.12717/DR.2024.28.1.13. PMid:38654977.

Martín-San Juan A, Martin-Hinojal CC, Nieto-Cristobal H, Martinez-Alborcia MJ, de Mercado E, Alvarez-Rodriguez M. Seminal extracellular vesicles from boar AI doses contain fertility-predictive protein and miRNA cargo and improve sperm physiology. bioRxiv. 2026. https://doi.org/10.64898/2026.03.16.712050.

Jung M, Rüdiger K, Schulze M. In vitro measures for assessing boar semen fertility. Reprod Domest Anim. 2015;50(S2 Suppl 2):20-4. https://doi.org/10.1111/rda.12533. PMid:26174915.

Kang S, Pang W-K, Ryu D-Y, Song W-H, Rahman MS, Park Y-J, Pang M-G. Porcine seminal protein-I and II mRNA expression in boar spermatozoa is significantly correlated with fertility. Theriogenology. 2019;138:31-8. https://doi.org/10.1016/j.theriogenology.2019.06.043. PMid:31280183.

Keller A, Maus M, Keller E, Kerns K. Deep learning classification method for boar sperm morphology analysis. Andrology. 2025;13(6):1615-25. https://doi.org/10.1111/andr.13758. PMid:39287620.

Khan MZ, Chen W, Naz S, Liu X, Liang H, Chen Y, Kou X, Liu Y, Ashraf I, Han Y, Peng Y, Wang C, Zahoor M. Determinant genetic markers of semen quality in livestock. Front Endocrinol (Lausanne). 2024;15:1456305. https://doi.org/10.3389/fendo.2024.1456305. PMid:39429738.

Kim K-U, Pang W-K, Kang S, Ryu D-Y, Song W-H, Rahman MS, Kwon W-S, Pang M-G. Sperm solute carrier family 9 regulator 1 is correlated with boar fertility. Theriogenology. 2019;126:254-60. https://doi.org/10.1016/j.theriogenology.2018.12.023. PMid:30590247.

Kim NH, Day BN, Lim JG, Lee HT, Chung KS. Effects of oviductal fluid and heparin on fertility and characteristics of porcine spermatozoa. Zygote. 1997;5(1):61-5. https://doi.org/10.1017/S0967199400003567. PMid:9223246.

Kiser JN, Clancey E, Moraes JGN, Dalton J, Burns GW, Spencer TE, Neibergs HL. Identification of loci associated with conception rate in primiparous Holstein cows. BMC Genomics. 2019;20(1):840. https://doi.org/10.1186/s12864-019-6203-2. PMid:31718557.

Knox RV. Artificial insemination in pigs today. Theriogenology. 2016;85(1):83-93. https://doi.org/10.1016/j.theriogenology.2015.07.009. PMid:26253434.

Kumaresan A, Das Gupta M, Datta TK, Morrell JM. Sperm DNA integrity and male fertility in farm animals: a review. Front Vet Sci. 2020;7:321. https://doi.org/10.3389/fvets.2020.00321. PMid:32637425.

Kwon W-S, Rahman MS, Lee J-S, Yoon S-J, Park Y-J, Pang M-G. Discovery of predictive biomarkers for litter size in boar spermatozoa. Mol Cell Proteomics. 2015;14(5):1230-40. https://doi.org/10.1074/mcp.M114.045369. PMid:25693803.

Kwon W-S, Rahman MS, Ryu D-Y, Khatun A, Pang M-G. Comparison of markers predicting litter size in different pig breeds. Andrology. 2017;5(3):568-77. https://doi.org/10.1111/andr.12332. PMid:28409901.

Kwon W-S, Shin D-H, Ryu D-Y, Khatun A, Rahman MS, Pang M-G. Applications of capacitation status for litter size enhancement in various pig breeds. Asian-Australas J Anim Sci. 2018;31(6):842-50. https://doi.org/10.5713/ajas.17.0760. PMid:29268576.

Langendijk P, Soede NM, Kemp B. Uterine activity, sperm transport, and the role of boar stimuli around insemination in sows. Theriogenology. 2005;63(2):500-13. https://doi.org/10.1016/j.theriogenology.2004.09.027. PMid:15626413.

Lee B-M, Park Y-J, Pang W-K, Ryu D-Y, Rahman MS, Lee D-Y, Pang M-G. Boar fertility is controlled through systematic changes of mitochondrial protein expression during sperm capacitation. Int J Biol Macromol. 2023;248:125955. https://doi.org/10.1016/j.ijbiomac.2023.125955. PMid:37494999.

Li B, Ming R, Qiu G, Guo H. Hypothalamic FTO-IGF2BP2-mediated m6A regulation of ANXA2: A novel axis preventing HS-induced sperm motility decline. Int J Biol Macromol. 2026;340(Pt 1):149223. https://doi.org/10.1016/j.ijbiomac.2025.149223. PMid:41360250.

Lian N, Niu Q, Lei Y, Li X, Li Y, Song X. MiR-221 is involved in depression by regulating Wnt2/CREB/BDNF axis in hippocampal neurons. Cell Cycle. 2018;17(24):2745-55. https://doi.org/10.1080/15384101.2018.1556060. PMid:30589396.

Lin C, Tholen E, Jennen D, Ponsuksili S, Schellander K, Wimmers K. Evidence for effects of testis and epididymis expressed genes on sperm quality and boar fertility traits. Reprod Domest Anim. 2006a;41(6):538-43. https://doi.org/10.1111/j.1439-0531.2006.00710.x. PMid:17107514.

Lin C-L, Jennen DGJ, Ponsuksili S, Tholen E, Tesfaye D, Schellander K, Wimmers K. Haplotype analysis of beta-actin gene for its association with sperm quality and boar fertility. J Anim Breed Genet. 2006b;123(6):384-8. https://doi.org/10.1111/j.1439-0388.2006.00622.x. PMid:17177693.

Lin Q, Cai X, Zhong Z, Li T, Chen X, Ayalew W, Xu Z, Wei C, Zhang X, Cheng H, Zhang Z, Li X, Tang Y, Chen S, Zhou J, Si J, Wu X, Ning C, Wang Q, Pan Y, Gao Y, Li J, Yu Y, Zhang Z, Zhao Y, Fang L, Zhang Z. Meta-GWAS of pig semen quality traits reveals conserved genes regulating mammalian fertility. Adv Sci (Weinh). 2026;13(16):e15203. https://doi.org/10.1002/advs.202515203. PMid:41504393.

Llavanera M. Evaluation of sperm quality and male fertility: the use of molecular markers in boar sperm and seminal plasma. Anim Reprod Sci. 2024;269:107545. https://doi.org/10.1016/j.anireprosci.2024.107545. PMid:38960838.

Llavanera M, Mateo-Otero Y, Viñolas-Vergés E, Bonet S, Yeste M. Sperm function, mitochondrial activity and in vivo fertility are associated to their mitochondrial DNA content in pigs. J Anim Sci Biotechnol. 2024;15(1):10. https://doi.org/10.1186/s40104-023-00988-0. PMid:38297401.

Lovercamp KW, Safranski TJ, Fischer KA, Manandhar G, Sutovsky M, Herring W, Sutovsky P. High resolution light microscopic evaluation of boar semen quality sperm cytoplasmic droplet retention in relationship with boar fertility parameters. Arch Androl. 2007a;53(4):219-28. https://doi.org/10.1080/01485010701426463. PMid:17852046.

Lovercamp KW, Safranski TJ, Fischer KA, Manandhar G, Sutovsky M, Herring W, Sutovsky P. Arachidonate 15-lipoxygenase and ubiquitin as fertility markers in boars. Theriogenology. 2007b;67(4):704-18. https://doi.org/10.1016/j.theriogenology.2006.08.019. PMid:17116325.

Martinez CA, Roca J, Alvarez-Rodriguez M, Rodriguez-Martinez H. miRNA-profiling in ejaculated and epididymal pig spermatozoa and their relation to fertility after artificial insemination. Biology (Basel). 2022;11(2):236. https://doi.org/10.3390/biology11020236. PMid:35205102.

Martínez-Hernández J, Garriga F, Ahmad A, Padilla L, Maside C, Bonet S, Barranco I, Roca J, Pastor LM, Yeste M. The oxytocin receptor in spermatozoa may originate from both spermatogenesis and epididymal maturation, and regulates capacitation. Andrology. 2026;14(3):747-65. https://doi.org/10.1111/andr.70123. PMid:41014065.

Martinez-Pastor F, Mata-Campuzano M, Alvarez-Rodriguez M, Alvarez M, Anel L, de Paz P. Probes and techniques for sperm evaluation by flow cytometry. Reprod Domest Anim. 2010;45(s2 Suppl 2):67-78. https://doi.org/10.1111/j.1439-0531.2010.01622.x. PMid:20591067.

Maside C, Recuero S, Salas-Huetos A, Ribas-Maynou J, Yeste M. Animal board invited review: an update on the methods for semen quality evaluation in swine - from farm to the lab. Animal. 2023;17(3):100720. https://doi.org/10.1016/j.animal.2023.100720. PMid:36801527.

Matás C, Martínez E, Vázquez JM, Roca J, Gadea J. In vitro penetration assay of boar sperm fertility: effect of various factors on the penetrability of immature pig oocytes. Theriogenology. 1996;46(3):503-13. https://doi.org/10.1016/0093-691X(96)00172-0. PMid:16727918.

Mateo-Otero Y. Integrating metabolomics into reproduction: sperm metabolism and fertility enhancement in pigs. Anim Reprod Sci. 2024;269:107539. https://doi.org/10.1016/j.anireprosci.2024.107539. PMid:38926002.

Mateo-Otero Y, Fernández-López P, Delgado-Bermúdez A, Nolis P, Roca J, Miró J, Barranco I, Yeste M. Metabolomic fingerprinting of pig seminal plasma identifies in vivo fertility biomarkers. J Anim Sci Biotechnol. 2021;12(1):113. https://doi.org/10.1186/s40104-021-00636-5. PMid:34772452.

Mateo-Otero Y, Llavanera M, Recuero S, Delgado-Bermúdez A, Barranco I, Ribas-Maynou J, Yeste M. Sperm DNA damage compromises embryo development, but not oocyte fertilisation in pigs. Biol Res. 2022a;55(1):15. https://doi.org/10.1186/s40659-022-00386-2. PMid:35365220.

Mateo-Otero Y, Ribas-Maynou J, Delgado-Bermúdez A, Llavanera M, Recuero S, Barranco I, Yeste M. Aldose reductase B1 in pig sperm is related to their function and fertilizing ability. Front Endocrinol. 2022b;13:773249. https://doi.org/10.3389/fendo.2022.773249. PMid:35173684.

Michailov Y, Ickowicz D, Breitbart H. Zn2+-stimulation of sperm capacitation and of the acrosome reaction is mediated by EGFR activation. Dev Biol. 2014;396(2):246-55. https://doi.org/10.1016/j.ydbio.2014.10.009. PMid:25446533.

Mills KM, Minton AM, Magee JM, Long JA. Pilot evaluation of sperm mobility for boar fertility classification using machine learning. Transl Anim Sci. 2026;10:txaf122. https://doi.org/10.1093/tas/txaf122. PMid:41799844.

Miyado K, Yoshida K, Yamagata K, Sakakibara K, Okabe M, Wang X, Miyamoto K, Akutsu H, Kondo T, Takahashi Y, Ban T, Ito C, Toshimori K, Nakamura A, Ito M, Miyado M, Mekada E, Umezawa A. The fusing ability of sperm is bestowed by CD9-containing vesicles released from eggs in mice. Proc Natl Acad Sci USA. 2008;105(35):12921-6. https://doi.org/10.1073/pnas.0710608105. PMid:18728192.

Myromslien FD, Tremoen NH, Andersen-Ranberg I, Fransplass R, Stenseth E-B, Zeremichael TT, van Son M, Grindflek E, Gaustad AH. Sperm DNA integrity in Landrace and Duroc boar semen and its relationship to litter size. Reprod Domest Anim. 2019;54(2):160-6. https://doi.org/10.1111/rda.13322. PMid:30168871.

Novak S, Ruiz-Sánchez A, Dixon WT, Foxcroft GR, Dyck MK. Seminal plasma proteins as potential markers of relative fertility in boars. J Androl. 2010;31(2):188-200. https://doi.org/10.2164/jandrol.109.007583. PMid:19713565.

Oh S-A, You Y-A, Park Y-J, Pang M-G. The sperm penetration assay predicts the litter size in pigs. Int J Androl. 2010;33(4):604-12. https://doi.org/10.1111/j.1365-2605.2009.00976.x. PMid:19538520.

Oliver SG, Winson MK, Kell DB, Baganz F. Systematic functional analysis of the yeast genome. Trends Biotechnol. 1998;16(9):373-8. https://doi.org/10.1016/S0167-7799(98)01214-1. PMid:9744112.

Padilla L, López-Arjona M, Martinez-Subiela S, Rodriguez-Martinez H, Roca J, Barranco I. Oxytocin in pig seminal plasma is positively related with in vivo fertility of inseminated sows. J Anim Sci Biotechnol. 2021;12(1):101. https://doi.org/10.1186/s40104-021-00620-z. PMid:34511116.

Pang W-K, Park Y-J, Pang M-G. Development of a biomolecular approach to identify sperm functions and fertility using sperm RNAs. Front Cell Dev Biol. 2023;11:1308167. https://doi.org/10.3389/fcell.2023.1308167. PMid:38116206.

Park M, Yoon H, Kang BH, Lee H, An J, Lee T, Cheong H-T, Lee S-H. Deep learning-based precision analysis for acrosome reaction by modification of plasma membrane in boar sperm. Animals (Basel). 2023a;13(16):2622. https://doi.org/10.3390/ani13162622. PMid:37627413.

Park Y-J, Pang W-K, Pang M-G. Integration of omics studies indicates that species-dependent molecular mechanisms govern male fertility. J Anim Sci Biotechnol. 2023b;14(1):28. https://doi.org/10.1186/s40104-023-00836-1. PMid:36859388.

Parra A, Martínez-Díaz P, Botía M, López-Arjona M, Lucas X, Barranco I, Roca J. The extracellular vesicles of boar seminal plasma contain oxytocin at levels associated with fertility. Extracell Vesicles Circ Nucleic Acids. 2025;6(4):1000-14. https://doi.org/10.20517/evcna.2025.94. PMid:41551609.

Pérez-Patiño C, Parrilla I, Barranco I, Vergara-Barberán M, Simó-Alfonso EF, Herrero-Martínez JM, Rodriguez-Martínez H, Martínez EA, Roca J. New in-depth analytical approach of the porcine seminal plasma proteome reveals potential fertility biomarkers. J Proteome Res. 2018;17(3):1065-76. https://doi.org/10.1021/acs.jproteome.7b00728. PMid:29411616.

Pérez-Patiño C, Parrilla I, Li J, Barranco I, Martínez EA, Rodriguez-Martínez H, Roca J. The proteome of pig spermatozoa is remodeled during ejaculation* [S]. Mol Cell Proteomics. 2019;18(1):41-50. https://doi.org/10.1074/mcp.RA118.000840. PMid:30257877.

Pértille F, Alvarez-Rodriguez M, da Silva AN, Barranco I, Roca J, Guerrero-Bosagna C, Rodriguez-Martinez H. Sperm methylome profiling can discern fertility levels in the porcine biomedical model. Int J Mol Sci. 2021;22(5):2679. https://doi.org/10.3390/ijms22052679. PMid:33800945.

Qiu T, Wang K, Li X, Jin J. MiR-671-5p inhibits gastric cancer cell proliferatioand promotes cell apoptosis by targeting URGCP. Exp Ther Med. 2018;16(6):4753-8. https://doi.org/10.3892/etm.2018.6813. PMid:30546398.

Reicks DL, Levis DG. Fertility of semen used in commercial production and the impact of sperm numbers and bacterial counts. Theriogenology. 2008;70(8):1377-9. https://doi.org/10.1016/j.theriogenology.2008.07.019. PMid:18768216.

Ren D, Xia J. Calcium signaling through CatSper channels in mammalian fertilization. Physiology (Bethesda). 2010;25(3):165-75. https://doi.org/10.1152/physiol.00049.2009. PMid:20551230.

Rex AS, Aagaard J, Fedder J. DNA fragmentation in spermatozoa: a historical review. Andrology. 2017;5(4):622-30. https://doi.org/10.1111/andr.12381. PMid:28718529.

Robertson SA, Redman CW, Mccracken SA, Hunt JS, Dimitriadis E, Moffett-King A, Chamley L. Immune modulators of implantation and placental development--a workshop report. Placenta. 2003;24(Suppl A):S16-20. https://doi.org/10.1053/plac.2002.0937. PMid:12842409.

Robertson SA, Sharkey DJ. The role of semen in induction of maternal immune tolerance to pregnancy. Semin Immunol. 2001;13(4):243-54. https://doi.org/10.1006/smim.2000.0320. PMid:11437632.

Roca J, Broekhuijse MLWJ, Parrilla I, Rodriguez‐Martinez H, Martinez EA, Bolarin A. Boar differences in artificial insemination outcomes: can they be minimized? Reprod Domest Anim. 2015;50(S2 Suppl 2):48-55. https://doi.org/10.1111/rda.12530. PMid:26174919.

Rodriguez I, Keller A, Jennett L, Johnson M, Shofner I, Mahmood M, Redel B, Kerns K. Capacitation-induced zinc ion flux and sperm plasma membrane remodeling predict porcine in vitro fertilization cleavage success. Mol Reprod Dev. 2026;93(1):e70085. https://doi.org/10.1002/mrd.70085. PMid:41560536.

Rodriguez-Caro H, Dragovic R, Shen M, Dombi E, Mounce G, Field K, Meadows J, Turner K, Lunn D, Child T, Southcombe JH, Granne I. In vitro decidualisation of human endometrial stromal cells is enhanced by seminal fluid extracellular vesicles. J Extracell Vesicles. 2019;8(1):1565262. https://doi.org/10.1080/20013078.2019.1565262. PMid:30728921.

Rodriguez-Martinez H. Semen evaluation and handling: Emerging techniques and future development. In: Chenoweth P, Lorton S, editors. Animal andrology: theories and applications. Wallingford: CABI; 2014. p. 509-49. https://doi.org/10.1079/9781780643168.0509.

Rodriguez-Martinez H. Semen evaluation techniques and their relationship with fertility. Anim Reprod. 2013;46:148-59. https://doi.org/10.1079/9781780643168.0509.

Rodriguez-Martinez H, Kvist U, Saravia F, Wallgren M, Johannisson A, Sanz L, Pena FJ, Martinez EA, Roca J, Vazquez JM, Calvete JJ. The physiological roles of the boar ejaculate. Soc Reprod Fertil Suppl. 2009;66:1-21. PMid:19848263.

Rodriguez-Martinez H, Martinez-Serrano CA, Alvarez-Rodriguez M, Martinez EA, Roca J. Reproductive physiology of the boar: what defines the potential fertility of an ejaculate? Anim Reprod Sci. 2024;269:107476. https://doi.org/10.1016/j.anireprosci.2024.107476. PMid:38664134.

Rodriguez-Martinez H, Nicander L, Viring S, Einarsson S, Larsson K. Ultrastructure of the uterotubal junction in preovulatory pigs. Anat Histol Embryol. 1990;19(1):16-36. https://doi.org/10.1111/j.1439-0264.1990.tb00875.x. PMid:2375508.

Rodriguez-Martinez H, Roca J. Extracellular vesicles in seminal plasma: A safe and relevant tool to improve fertility in livestock? Anim Reprod Sci. 2022;244:107051. https://doi.org/10.1016/j.anireprosci.2022.107051. PMid:35933831.

Rodriguez-Martinez H, Roca J, Alvarez-Rodriguez M, Martinez-Serrano CA. How does the boar epididymis regulate the emission of fertile spermatozoa? Anim Reprod Sci. 2022;246:106829. https://doi.org/10.1016/j.anireprosci.2021.106829. PMid:34452796.

Roudebush WE, Diehl JR. Platelet-activating factor content in boar spermatozoa correlates with fertility. Theriogenology. 2001;55(8):1633-8. https://doi.org/10.1016/S0093-691X(01)00508-8. PMid:11396471.

Rozeboom KJ, Troedsson MH, Hodson HH, Shurson GC, Crabo BG. The importance of seminal plasma on the fertility of subsequent artificial inseminations in swine. J Anim Sci. 2000;78(2):443-8. https://doi.org/10.2527/2000.782443x. PMid:10709936.

Ruiz-Conca M, Gardela J, Martínez CA, Wright D, López-Bejar M, Rodríguez-Martínez H, Álvarez-Rodríguez M. Natural mating differentially triggers expression of glucocorticoid receptor (NR3C1)-related genes in the preovulatory porcine female reproductive tract. Int J Mol Sci. 2020;21(12):4437. https://doi.org/10.3390/ijms21124437. PMid:32580389.

Ryu DY, Pang WK, Rahman MS, Park YJ, Pang MG. Peroxiredoxin 4 directly affects the male fertility outcome in porcine. Theriogenology. 2021;171:85-93. https://doi.org/10.1016/j.theriogenology.2021.05.020. PMid:34051589.

Sá P, Gòdia M, Godinho RM, Sevillano CA, Harlizius B, Madsen O, Bovenhuis H. A genome-wide association study reveals additive and recessive alleles affecting male fertility in pigs. J Anim Sci Biotechnol. 2025a;16(1):171. https://doi.org/10.1186/s40104-025-01312-8. PMid:41392145.

Sá P, Godinho RM, Gòdia M, Sevillano CA, Harlizius B, Madsen O, Bovenhuis H. Genetic parameters and parental and early-life effects of boar semen traits. Genet Sel Evol. 2025b;57(1):4. https://doi.org/10.1186/s12711-025-00954-6. PMid:39915704.

Schulze M, Ruediger K, Mueller K, Jung M, Well C, Reissmann M. Development of an in vitro index to characterize fertilizing capacity of boar ejaculates. Anim Reprod Sci. 2013;140(1-2):70-6. https://doi.org/10.1016/j.anireprosci.2013.05.012. PMid:23773327.

Sellés E, Gadea J, Romar R, Matás C, Ruiz S. Analysis of in vitro fertilizing capacity to evaluate the freezing procedures of boar semen and to predict the subsequent fertility. Reprod Domest Anim. 2003;38(1):66-72. https://doi.org/10.1046/j.1439-0531.2003.00406.x. PMid:12535333.

Sonderman JP, Luebbe JJ. Semen production and fertility issues related to differences in genetic lines of boars. Theriogenology. 2008;70(8):1380-3. https://doi.org/10.1016/j.theriogenology.2008.08.009. PMid:18783820.

Sutkeviciene N, Riskeviciene V, Januskauskas A, Zilinskas H, Andersson M. Assessment of sperm quality traits in relation to fertility in boar semen. Acta Vet Scand. 2009;51(1):53. https://doi.org/10.1186/1751-0147-51-53. PMid:20015377.

Sutovsky P, Hamilton LE, Zigo M, Ortiz D’Avila Assumpção ME, Jones A, Tirpak F, Agca Y, Kerns K, Sutovsky M. Biomarker-based human and animal sperm phenotyping: the good, the bad and the ugly†. Biol Reprod. 2024;110(6):1135-56. https://doi.org/10.1093/biolre/ioae061. PMid:38640912.

Tardif S, Laforest JP, Cormier N, Bailey JL. The importance of porcine sperm parameters on fertility in vivo. Theriogenology. 1999;52(3):447-59. https://doi.org/10.1016/S0093-691X(99)00142-9. PMid:10734379.

Tjalsma H, Bolhuis A, Jongbloed JD, Bron S, van Dijl JM. Signal peptide-dependent protein transport in Bacillus subtilis: a genome-based survey of the secretome. Microbiol Mol Biol Rev. 2000;64(3):515-47. https://doi.org/10.1128/MMBR.64.3.515-547.2000. PMid:10974125.

Toledo-Guardiola SM, Luongo C, Martínez-Pastor F, Soriano-Úbeda C, Matás C. The individual variations in sperm quality of high-fertility boars impact the offspring production and early physiological functions. Vet Sci. 2025;12(6):582. https://doi.org/10.3390/vetsci12060582. PMid:40559819.

Tremoen NH, Gaustad AH, Andersen-Ranberg I, van Son M, Zeremichael TT, Frydenlund K, Grindflek E, Våge DI, Myromslien FD. Relationship between sperm motility characteristics and ATP concentrations, and association with fertility in two different pig breeds. Anim Reprod Sci. 2018;193:226-34. https://doi.org/10.1016/j.anireprosci.2018.04.075. PMid:29703418.

Tsai PS, Gadella BM. Molecular kinetics of proteins at the surface of porcine sperm before and during fertilization. Soc Reprod Fertil Suppl. 2009;66:23-36. PMid:19848264.

Tsakmakidis IA, Lymberopoulos AG, Khalifa TA. Relationship between sperm quality traits and field-fertility of porcine semen. J Vet Sci. 2010;11(2):151-4. https://doi.org/10.4142/jvs.2010.11.2.151. PMid:20458156.

Vadnais ML, Roberts KP. Seminal plasma proteins inhibit in vitro- and cooling-induced capacitation in boar spermatozoa. Reprod Fertil Dev. 2010;22(6):893-900. https://doi.org/10.1071/RD09274. PMid:20591323.

Vicente-Carrillo A, Álvarez-Rodríguez M, Rodríguez-Martínez H. The CatSper channel modulates boar sperm motility during capacitation. Reprod Biol. 2017;17(1):69-78. https://doi.org/10.1016/j.repbio.2017.01.001. PMid:28077244.

Waberski D, Magnus F, Mendonca Ferreira F, Petrunkina AM, Weitze KF, Töpfer-Petersen E. Importance of sperm-binding assays for fertility prognosis of porcine spermatozoa. Theriogenology. 2005;63(2):470-84. https://doi.org/10.1016/j.theriogenology.2004.09.025. PMid:15626412.

Wang H, Liu J, Cho K-H, Ren D. A novel, single, transmembrane protein CATSPERG is associated with CATSPER1 channel protein. Biol Reprod. 2009;81(3):539-44. https://doi.org/10.1095/biolreprod.109.077107. PMid:19516020.

Wei X, Moncada-Pazos A, Cal S, Soria-Valles C, Gartner J, Rudloff U, Lin JC, Rosenberg SA, López-Otín C, Samuels Y. Analysis of the disintegrin-metalloproteinases family reveals ADAM29 and ADAM7 are often mutated in melanoma. Hum Mutat. 2011;32(6):E2148-75. https://doi.org/10.1002/humu.21477. PMid:21618342.

Xu Z, Xie Y, Wu C, Gu T, Zhang X, Yang J, Yang H, Zheng E, Huang S, Xu Z, Li Z, Cai G, Liu D, Hong L, Wu Z. Xu, Zheng, Li, Z., Cai, G., Liu, D., Hong, L., Wu, Z. The effects of boar seminal plasma extracellular vesicles on sperm fertility. Theriogenology. 2024;213:79-89. https://doi.org/10.1016/j.theriogenology.2023.09.026. PMid:37816296.

Yatsenko AN, Georgiadis AP, Murthy LJ, Lamb DJ, Matzuk MM. UBE2B mRNA alterations are associated with severe oligozoospermia in infertile men. Mol Hum Reprod. 2013;19(6):388-94. https://doi.org/10.1093/molehr/gat008. PMid:23378580.

Yeste M, Briz M, Pinart E, Sancho S, Bussalleu E, Bonet S. The osmotic tolerance of boar spermatozoa and its usefulness as sperm quality parameter. Anim Reprod Sci. 2010;119(3-4):265-74. https://doi.org/10.1016/j.anireprosci.2010.02.011. PMid:20227204.

Zeng F, Chen Y, Guo C, Li C, Wei H, Li L, Meng L, Zhang S. Analysis of differentially abundant proteins related to boar fertility in seminal plasma using iTRAQ-based quantitative proteomics. J Proteomics. 2021;236:104120. https://doi.org/10.1016/j.jprot.2021.104120. PMid:33540064.

Zhang J, Liu H, Yang Q, Li P, Wen Y, Han X, Li B, Jiang H, Li X. Genomic sequencing reveals the diversity of seminal bacteria and relationships to reproductive potential in boar sperm. Front Microbiol. 2020;11:1873. https://doi.org/10.3389/fmicb.2020.01873. PMid:32903829.

Zhang X, Yazaki J, Sundaresan A, Cokus S, Chan SW-L, Chen H, Henderson IR, Shinn P, Pellegrini M, Jacobsen SE, Ecker JR. Genome-wide high-resolution mapping and functional analysis of DNA methylation in arabidopsis. Cell. 2006;126(6):1189-201. https://doi.org/10.1016/j.cell.2006.08.003. PMid:16949657.

Zhang Y, Liu Y, Liang H, Xu Q, Liu Z, Weng X. Metabolomic differences of seminal plasma between boars with high and low average conception rates after artificial insemination. Reprod Domest Anim. 2021;56(1):161-71. https://doi.org/10.1111/rda.13861. PMid:33176019.
 


Submitted date:
03/18/2026

Accepted date:
08/06/2026

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