{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T22:06:30Z","timestamp":1792101990332,"version":"4.4.0"},"reference-count":78,"publisher":"MDPI AG","issue":"15","license":[{"start":{"date-parts":[[2025,8,5]],"date-time":"2025-08-05T00:00:00Z","timestamp":1754352000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Male infertility is a multifactorial condition often associated with disruptions in sperm metabolism and mitochondrial function, yet traditional semen analysis provides limited insight into these molecular mechanisms. Understanding sperm bioenergetics and metabolic dysfunctions is crucial for improving the diagnosis and treatment of conditions such as asthenozoospermia and azoospermia. This systematic review synthesizes recent literature, focusing on advanced tools and techniques\u2014including omics technologies, advanced imaging, spectroscopy, and functional assays\u2014that enable comprehensive molecular assessment of sperm metabolism and development. The reviewed studies highlight the effectiveness of metabolomics, proteomics, and transcriptomics in identifying metabolic biomarkers linked to male infertility. Non-invasive imaging modalities such as Raman and magnetic resonance spectroscopy offer real-time metabolic profiling, while the seminal microbiome is increasingly recognized for its role in modulating sperm metabolic health. Despite these advances, challenges remain in clinical validation and implementation of these techniques in routine infertility diagnostics. Integrating molecular metabolic assessments with conventional semen analysis promises enhanced diagnostic precision and personalized therapeutic approaches, ultimately improving reproductive outcomes. Continued research is needed to standardize biomarkers and validate clinical utility. Furthermore, these metabolic tools hold significant potential to elucidate the underlying causes of previously misunderstood and unexplained infertility cases, offering new avenues for diagnosis and treatment.<\/jats:p>","DOI":"10.3390\/ijms26157544","type":"journal-article","created":{"date-parts":[[2025,8,6]],"date-time":"2025-08-06T07:45:11Z","timestamp":1754466311000},"page":"7544","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":15,"title":["Exploring Human Sperm Metabolism and Male Infertility: A Systematic Review of Genomics, Proteomics, Metabolomics, and Imaging Techniques"],"prefix":"10.3390","volume":"26","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-1355-4633","authenticated-orcid":false,"given":"Achraf","family":"Zakaria","sequence":"first","affiliation":[{"name":"Research Laboratory of Microbiology, Infectious Diseases, Allergology and Pathogen Surveillance (LARMIAS), Mohammed VI Faculty of Medicine, Mohammed VI University of Sciences and Health (UM6SS), Casablanca 82403, Morocco"},{"name":"LABOMAC IVF Centers and Clinical Laboratory Medicine, Casablanca 20080, Morocco"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4122-2294","authenticated-orcid":false,"given":"Idrissa","family":"Diawara","sequence":"additional","affiliation":[{"name":"Research Laboratory of Microbiology, Infectious Diseases, Allergology and Pathogen Surveillance (LARMIAS), Mohammed VI Faculty of Medicine, Mohammed VI University of Sciences and Health (UM6SS), Casablanca 82403, Morocco"},{"name":"Mohammed VI Higher Institute of Biosciences and Biotechnologies, Mohammed VI University of Sciences and Health (UM6SS), Casablanca 82403, Morocco"},{"name":"Research Unit, Mohammed VI Center for Research and Innovation, Rabat 10112, Morocco"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amal","family":"Bouziyane","sequence":"additional","affiliation":[{"name":"Research Laboratory of Microbiology, Infectious Diseases, Allergology and Pathogen Surveillance (LARMIAS), Mohammed VI Faculty of Medicine, Mohammed VI University of Sciences and Health (UM6SS), Casablanca 82403, Morocco"},{"name":"Cheikh Khalifa International University Hospital, Casablanca 82403, Morocco"},{"name":"Mohammed VI International University Hospital, Bouskoura 27182, Morocco"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Noureddine","family":"Louanjli","sequence":"additional","affiliation":[{"name":"LABOMAC IVF Centers and Clinical Laboratory Medicine, Casablanca 20080, Morocco"},{"name":"IVF Center IRIFIV, Iris Clinic, Casablanca 20330, Morocco"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,8,5]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/0005-2728(86)90010-1","article-title":"Control of respiration and of motility in ejaculated bull spermatozoa","volume":"850","author":"Bohnensack","year":"1986","journal-title":"Biochim. Et Biophys. Acta (BBA)-Bioenerg."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1007\/s10815-014-0425-7","article-title":"Contemporary evidence on the physiological role of reactive oxygen species in human sperm function","volume":"32","author":"Agarwal","year":"2015","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Visconti, P. (2012). Sperm Bioenergetics in a Nutshell1. Biol. Reprod., 87.","DOI":"10.1095\/biolreprod.112.104109"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1111\/andr.13468","article-title":"Intracytoplasmic sperm injection vs. in-vitro fertilization in couples in whom the male partners had a semen analysis within normal reference ranges: An open debate","volume":"12","author":"Sciorio","year":"2023","journal-title":"Andrology"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"hoaf001","DOI":"10.1093\/hropen\/hoaf001","article-title":"Now is the time to introduce new innovative assisted reproduction methods to implement accessible, affordable, and demonstrably successful advanced infertility services in resource-poor countries","volume":"2025","author":"Ombelet","year":"2025","journal-title":"Hum. Reprod. Open"},{"key":"ref_6","unstructured":"World Health Organization (2025, April 17). Infertility. Available online: https:\/\/www.who.int\/news-room\/fact-sheets\/detail\/infertility."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Agarwal, A., Mulgund, A., Hamada, A., and Chyatte, M.R. (2015). A unique view on male infertility around the globe. Reprod. Biol. Endocrinol., 13.","DOI":"10.1186\/s12958-015-0032-1"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1074\/mcp.M112.020552","article-title":"Human Sperm Tail Proteome Suggests New Endogenous Metabolic Pathways","volume":"12","author":"Amaral","year":"2013","journal-title":"Mol. Cell Proteom."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"He, J., Ma, M., Xu, Z., Guo, J., Chen, H., Yang, X., Chen, P., and Liu, G. (2024). Association between semen microbiome disorder and sperm DNA damage. Microbiol. Spectr., 12.","DOI":"10.1128\/spectrum.00759-24"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"2813","DOI":"10.1210\/jc.2013-4148","article-title":"Novel Biomarkers of Androgen Deficiency From Seminal Plasma Profiling Using High-Resolution Mass Spectrometry","volume":"99","author":"Milardi","year":"2014","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Sharma, R., Agarwal, A., Mohanty, G., Jesudasan, R., Gopalan, B., Willard, B., Yadav, S.P., and Sabanegh, E. (2013). Functional proteomic analysis of seminal plasma proteins in men with various semen parameters. Reprod. Biol. Endocrinol., 11.","DOI":"10.1186\/1477-7827-11-38"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Sharma, R., Agarwal, A., Mohanty, G., Hamada, A.J., Gopalan, B., Willard, B., Yadav, S., and du Plessis, S. (2013). Proteomic analysis of human spermatozoa proteins with oxidative stress. Reprod. Biol. Endocrinol., 11.","DOI":"10.1186\/1477-7827-11-48"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"e23772","DOI":"10.1002\/mrd.23772","article-title":"Sperm Metabolism","volume":"91","author":"Abruzzese","year":"2024","journal-title":"Mol. Reprod. Dev."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1087","DOI":"10.1093\/humrep\/dey072","article-title":"Exosomal microRNAs in seminal plasma are markers of the origin of azoospermia and can predict the presence of sperm in testicular tissue","volume":"33","author":"Mata","year":"2018","journal-title":"Hum. Reprod."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Kyrgiafini, M.-A., Giannoulis, T., Chatziparasidou, A., Christoforidis, N., and Mamuris, Z. (2023). Unveiling the Genetic Complexity of Teratozoospermia: Integrated Genomic Analysis Reveals Novel Insights into lncRNAs\u2019 Role in Male Infertility. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms241915002"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1111\/andr.13461","article-title":"Seminal extracellular vesicle sncRNA sequencing reveals altered miRNA\/isomiR profiles as sperm retrieval biomarkers for azoospermia","volume":"12","author":"Larriba","year":"2023","journal-title":"Andrology"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"951","DOI":"10.2217\/epi-2022-0122","article-title":"Genome-Wide Methylation Analyses of Human Sperm Unravel Novel Differentially Methylated Regions in Asthenozoospermia","volume":"14","author":"Li","year":"2022","journal-title":"Epigenomics"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"687","DOI":"10.1111\/andr.13394","article-title":"MicroRNAs expression in semen and testis of azoospermic men","volume":"11","author":"Wainstein","year":"2023","journal-title":"Andrology"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1722","DOI":"10.1373\/clinchem.2011.169714","article-title":"Altered Profile of Seminal Plasma MicroRNAs in the Molecular Diagnosis of Male Infertility","volume":"57","author":"Wang","year":"2011","journal-title":"Clin. Chem."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1186\/s12014-015-9094-5","article-title":"Comparative proteomic network signatures in seminal plasma of infertile men as a function of reactive oxygen species","volume":"12","author":"Agarwal","year":"2015","journal-title":"Clin. Proteom."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"707","DOI":"10.4103\/1008-682X.185848","article-title":"AMP-activated kinase in human spermatozoa: Identification, intracellular localization, and key function in the regulation of sperm motility","volume":"19","author":"Bragado","year":"2017","journal-title":"Asian J. Androl."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1007\/s11306-024-02118-x","article-title":"Lipidomics random forest algorithm of seminal plasma is a promising method for enhancing the diagnosis of necrozoospermia","volume":"20","author":"Deng","year":"2024","journal-title":"Metabolomics"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Fietz, D., Sgaier, R., O\u2019donnell, L., Stanton, P.G., Dagley, L.F., Webb, A.I., Schuppe, H.-C., Diemer, T., and Pilatz, A. (2024). Proteomic biomarkers in seminal plasma as predictors of reproductive potential in azoospermic men. Front. Endocrinol., 15.","DOI":"10.3389\/fendo.2024.1327800"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"853","DOI":"10.1007\/s10815-015-0465-7","article-title":"A proteomic analysis on human sperm tail: Comparison between normozoospermia and asthenozoospermia","volume":"32","author":"Hashemitabar","year":"2015","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"929","DOI":"10.1007\/s10930-021-10013-w","article-title":"Proteomic Profile of Sperm in Infertile Males Reveals Changes in Metabolic Pathways","volume":"40","author":"Liang","year":"2021","journal-title":"Protein J."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1192","DOI":"10.1111\/andr.12991","article-title":"Altered mitochondrial function in spermatozoa from patients with repetitive fertilization failure after ICSI revealed by proteomics","volume":"9","author":"Jodar","year":"2021","journal-title":"Andrology"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Martins, A.D., Selvam, M.K.P., Agarwal, A., Alves, M.G., and Baskaran, S. (2020). Alterations in seminal plasma proteomic profile in men with primary and secondary infertility. Sci. Rep., 10.","DOI":"10.1038\/s41598-020-64434-1"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1998","DOI":"10.1074\/mcp.M116.058289","article-title":"Intact Cell MALDI-TOF MS on Sperm: A Molecular Test For Male Fertility Diagnosis","volume":"15","author":"Soler","year":"2016","journal-title":"Mol. Cell Proteom."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Sulc, A., Cz\u00e9t\u00e1ny, P., M\u00e1t\u00e9, G., Ball\u00f3, A., Semj\u00e9n, D., Sz\u00e1nt\u00f3, \u00c1., and M\u00e1rk, L. (2024). MALDI Imaging Mass Spectrometry Reveals Lipid Alterations in Physiological and Sertoli Cell-Only Syndrome Human Testicular Tissue Sections. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25158358"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1530\/REP-17-0746","article-title":"RNASET2 impairs the sperm motility via PKA\/PI3K\/calcium signal pathways","volume":"155","author":"Xu","year":"2018","journal-title":"Reproduction"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1859","DOI":"10.1111\/andr.12882","article-title":"Metabolomic signature of the seminal plasma in men with severe oligoasthenospermia","volume":"8","author":"Boguenet","year":"2020","journal-title":"Andrology"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1093\/molehr\/gay046","article-title":"Probing human Kyrgiafini using 13C-magnetic resonance spectroscopy","volume":"25","author":"Calvert","year":"2018","journal-title":"Mol. Hum. Reprod."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1007\/s10815-014-0282-4","article-title":"Identification of biochemical differences between different forms of male infertility by nuclear magnetic resonance (NMR) spectroscopy","volume":"31","author":"Jayaraman","year":"2014","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1007\/s11306-023-02020-y","article-title":"Low-polarity untargeted metabolomic profiling as a tool to gain insight into seminal fluid","volume":"19","author":"Olesti","year":"2023","journal-title":"Metabolomics"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"121929","DOI":"10.1016\/j.jchromb.2019.121929","article-title":"Steroid profile analysis by LC-HRMS in human seminal fluid","volume":"1136","author":"Olesti","year":"2020","journal-title":"J. Chromatogr. B"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1111\/andr.12027","article-title":"Identification of endogenous metabolites in human sperm cells using proton nuclear magnetic resonance (1H-NMR) spectroscopy and gas chromatography-mass spectrometry (GC-MS)","volume":"3","author":"Paiva","year":"2015","journal-title":"Andrology"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Qiao, S., Wu, W., Chen, M., Tang, Q., Xia, Y., Jia, W., and Wang, X. (2017). Seminal plasma metabolomics approach for the diagnosis of unexplained male infertility. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0181115"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1093\/molehr\/gax025","article-title":"1H Magnetic Resonance Spectroscopy of live human sperm","volume":"23","author":"Reynolds","year":"2017","journal-title":"Mol. Hum. Reprod."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Zhao, K., Zhang, J., Xu, Z., Xu, Y., Xu, A., Chen, W., Miao, C., Liu, S., Wang, Z., and Jia, R. (2018). Metabolomic Profiling of Human Spermatozoa in Idiopathic Asthenozoospermia Patients Using Gas Chromatography-Mass Spectrometry. BioMed Res. Int., 2018.","DOI":"10.1155\/2018\/8327506"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Zhao, K., Zhang, Q., Cong, R., Xu, Z., Xu, Y., and Han, J. (2024). Metabolomic profiling of human semen in patients with oligospermia using high performance liquid chromatography-tandem mass spectrometry. Sci. Rep., 14.","DOI":"10.1038\/s41598-024-74658-0"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"e407","DOI":"10.1111\/j.1365-2605.2011.01151.x","article-title":"Caspase-3 detection in human testicular spermatozoa from azoospermic and non-azoospermic patients","volume":"34","author":"Almeida","year":"2011","journal-title":"Int. J. Androl."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Cassina, A., Silveira, P., Cantu, L., Montes, J.M., Radi, R., and Sapiro, R. (2015). Defective Human Sperm Cells Are Associated with Mitochondrial Dysfunction and Oxidant Production1. Biol. Reprod., 93.","DOI":"10.1095\/biolreprod.115.130989"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Irigoyen, P., Pintos-Polasky, P., Rosa-Villagran, L., Skowronek, M.F., Cassina, A., and Sapiro, R. (2022). Mitochondrial metabolism determines the functional status of human sperm and correlates with semen parameters. Front. Cell Dev. Biol., 10.","DOI":"10.3389\/fcell.2022.926684"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1188","DOI":"10.1111\/and.12558","article-title":"Seminal level of clusterin in infertile men as a significant biomarker reflecting spermatogenesis","volume":"48","author":"Fukuda","year":"2016","journal-title":"Andrologia"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"2315","DOI":"10.1016\/j.fertnstert.2011.03.059","article-title":"Mitochondrial membrane potential profile and its correlation with increasing sperm motility","volume":"95","author":"Paoli","year":"2011","journal-title":"Fertil. Steril."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Da Costa, R., Amaral, S., Redmann, K., Kliesch, S., and Schlatt, S. (2018). Spectral features of nuclear DNA in human sperm assessed by Raman Microspectroscopy: Effects of UV-irradiation and hydration. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0207786"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1016\/j.fertnstert.2014.03.035","article-title":"Raman spectroscopy as an ex vivo noninvasive approach to distinguish complete and incomplete spermatogenesis within human seminiferous tubules","volume":"102","author":"Liu","year":"2014","journal-title":"Fertil. Steril."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"2604","DOI":"10.1016\/j.fertnstert.2010.03.022","article-title":"Comparison of chemiluminescence and flow cytometry in the estimation of reactive oxygen and nitrogen species in human semen","volume":"94","author":"Aziz","year":"2010","journal-title":"Fertil. Steril."},{"key":"ref_49","first-page":"293","article-title":"[Ca2+]i oscillations in human sperm are triggered in the flagellum by membrane potential-sensitive activity of CatSper","volume":"36","author":"Brown","year":"2020","journal-title":"Hum. Reprod."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1307","DOI":"10.3390\/cimb43030093","article-title":"A Comparative Analysis of the Altered Levels of Human Seminal Plasma Constituents as Contributing Factors in Different Types of Male Infertility","volume":"43","author":"Vashisht","year":"2021","journal-title":"Curr. Issues Mol. Biol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1016\/j.redox.2015.11.010","article-title":"Interplay between oxidant species and energy metabolism","volume":"8","author":"Quijano","year":"2016","journal-title":"Redox Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1016\/0092-8674(85)90085-6","article-title":"Metabolite channeling: A phosphorylcreatine shuttle to mediate high energy phosphate transport between sperm mitochondrion and tail","volume":"41","author":"Tombes","year":"1985","journal-title":"Cell"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1093\/humupd\/dmi053","article-title":"Glycolysis and sperm motility: Does a spoonful of sugar help the flagellum go round?","volume":"12","author":"Ford","year":"2006","journal-title":"Hum. Reprod. Update"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1038\/157079a0","article-title":"Fructose, a Constituent of Semen","volume":"157","author":"Mann","year":"1946","journal-title":"Nature"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1586","DOI":"10.1095\/biolreprod.103.019216","article-title":"Aldose Reductase and Macrophage Migration Inhibitory Factor Are Associated with Epididymosomes and Spermatozoa in the Bovine Epididymis1","volume":"69","author":"Frenette","year":"2003","journal-title":"Biol. Reprod."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1530\/rep.0.1230579","article-title":"Differential effects of glucose and fructose on hexose metabolism in dog spermatozoa","volume":"123","author":"Rigau","year":"2002","journal-title":"Reproduction"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"14523","DOI":"10.1016\/S0021-9258(18)42067-4","article-title":"Fructose transporter in human spermatozoa and small intestine is GLUT5","volume":"267","author":"Burant","year":"1992","journal-title":"J. Biol. Chem."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"348","DOI":"10.2164\/jandrol.110.011197","article-title":"GLUTs and Mammalian Sperm Metabolism","volume":"32","author":"Bucci","year":"2010","journal-title":"J. Androl."},{"key":"ref_59","first-page":"226","article-title":"Sorbitol metabolism in spermatozoa","volume":"151","author":"King","year":"1959","journal-title":"Proc. R. Soc. Lond. Ser. B Biol. Sci."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1095\/biolreprod.108.068882","article-title":"Sorbitol Can Fuel Mouse Sperm Motility and Protein Tyrosine Phosphorylation via Sorbitol Dehydrogenase1","volume":"80","author":"Cao","year":"2009","journal-title":"Biol. Reprod."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Darr, C.R., Varner, D.D., Teague, S., Cortopassi, G.A., Datta, S., and Meyers, S.A. (2016). Lactate and Pyruvate Are Major Sources of Energy for Stallion Sperm with Dose Effects on Mitochondrial Function, Motility, and ROS Production. Biol. Reprod., 95.","DOI":"10.1095\/biolreprod.116.140707"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1042\/bj3430281","article-title":"The proton-linked monocarboxylate transporter (MCT) family: Structure, function and regulation","volume":"343","author":"Halestrap","year":"1999","journal-title":"Biochem. J."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"694","DOI":"10.4103\/1008-682X.124561","article-title":"The genetic variation in Monocarboxylic acid transporter 2 (MCT2) has functional and clinical relevance with male infertility","volume":"16","author":"Lee","year":"2014","journal-title":"Asian J. Androl."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1126\/science.139.3555.601","article-title":"Lactate Dehydrogenases in Human Testes","volume":"139","author":"Blanco","year":"1963","journal-title":"Science"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1530\/REP-14-0500","article-title":"Investigation of the stallion sperm proteome by mass spectrometry","volume":"149","author":"Swegen","year":"2015","journal-title":"Reproduction"},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Zhu, Z., Li, R., Feng, C., Liu, R., Zheng, Y., Hoque, S.A.M., Wu, D., Lu, H., Zhang, T., and Zeng, W. (2020). Exogenous Oleic Acid and Palmitic Acid Improve Boar Sperm Motility via Enhancing Mitochondrial \u0392-Oxidation for ATP Generation. Animals, 10.","DOI":"10.3390\/ani10040591"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1002\/rmb2.12381","article-title":"Saturated fatty acids accelerate linear motility through mitochondrial ATP production in bull sperm","volume":"20","author":"Islam","year":"2021","journal-title":"Reprod. Med. Biol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"R163","DOI":"10.1530\/REP-13-0178","article-title":"Mitochondria functionality and sperm quality","volume":"146","author":"Amaral","year":"2013","journal-title":"Reproduction"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1002\/(SICI)1098-2795(200006)56:2<207::AID-MRD12>3.0.CO;2-4","article-title":"Evidence for a functional glycogen metabolism in mature mammalian spermatozoa","volume":"56","author":"Ballester","year":"2000","journal-title":"Mol. Reprod. Dev."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1002\/mrd.10268","article-title":"Glucose- and fructose-induced dog-sperm glycogen synthesis shows specific changes in the location of the sperm glycogen deposition","volume":"64","author":"Palomo","year":"2003","journal-title":"Mol. Reprod. Dev."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/rda.12901","article-title":"Effect of mitochondrial uncoupling and glycolysis inhibition on ram sperm functionality","volume":"52","author":"Losano","year":"2017","journal-title":"Reprod. Domest. Anim."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"106779","DOI":"10.1016\/j.anireprosci.2021.106779","article-title":"Effect of glucose concentration and cryopreservation on mitochondrial functions of bull spermatozoa and relationship with sire conception rate","volume":"230","author":"Moraes","year":"2021","journal-title":"Anim. Reprod. Sci."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.theriogenology.2021.09.015","article-title":"Effects from disruption of mitochondrial electron transport chain function on bull sperm motility","volume":"176","author":"Bulkeley","year":"2021","journal-title":"Theriogenology"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Balbach, M., Ghanem, L., Violante, S., Kyaw, A., Romarowski, A., Cross, J.R., Visconti, P.E., Levin, L.R., and Buck, J. (2023). Capacitation induces changes in metabolic pathways supporting motility of epididymal and ejaculated sperm. Front. Cell Dev. Biol., 11.","DOI":"10.3389\/fcell.2023.1160154"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"3600","DOI":"10.18632\/oncotarget.12274","article-title":"Factors and pathways involved in capacitation: How are they regulated?","volume":"8","author":"Jin","year":"2016","journal-title":"Oncotarget"},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Tourmente, M., Sansegundo, E., and Roldan, E.R.S. (2022). Capacitation promotes a shift in energy metabolism in murine sperm. Front. Cell Dev. Biol., 10.","DOI":"10.3389\/fcell.2022.950979"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Mateo-Otero, Y., Madrid-Gambin, F., Llavanera, M., Gomez-Gomez, A., Haro, N., Pozo, O.J., and Yeste, M. (2023). Sperm physiology and in vitro fertilising ability rely on basal metabolic activity: Insights from the pig model. Commun. Biol., 6.","DOI":"10.1038\/s42003-023-04715-3"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Li, L., Hao, X., Chen, H., Wang, L., Chen, A., Song, X., Hu, Z., Su, Y., Lin, H., and Fan, P. (2020). Metabolomic characterization of semen from asthenozoospermic patients using ultra-high-performance liquid chromatography\u2013tandem quadrupole time-of-flight mass spectrometry. Biomed. Chromatogr., 34.","DOI":"10.1002\/bmc.4897"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/15\/7544\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T18:23:14Z","timestamp":1760034194000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/15\/7544"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,5]]},"references-count":78,"journal-issue":{"issue":"15","published-online":{"date-parts":[[2025,8]]}},"alternative-id":["ijms26157544"],"URL":"https:\/\/doi.org\/10.3390\/ijms26157544","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,5]]}}}