{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T10:16:29Z","timestamp":1788430589743,"version":"build-2803163510"},"reference-count":70,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T00:00:00Z","timestamp":1788307200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Guangdong S&T Programme","award":["2024B1212060001"],"award-info":[{"award-number":["2024B1212060001"]}]},{"name":"Southern Marine Science and Engineering Guangdong Laboratory","award":["SML2024SP001"],"award-info":[{"award-number":["SML2024SP001"]}]},{"award":["SML2024SP001"],"award-info":[{"award-number":["SML2024SP001"]}],"id":[{"id":"https:\/\/ror.org\/03swgqh13","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012245","name":"Science and Technology Planning Project of Guangdong Province","doi-asserted-by":"publisher","award":["2023B1212060023"],"award-info":[{"award-number":["2023B1212060023"]}],"id":[{"id":"10.13039\/501100012245","id-type":"DOI","asserted-by":"publisher"}]},{"name":"2025 Guangdong Provincial Dedicated Funds for State Key Laboratories, Agriculture Research System of China","award":["ARS-47"],"award-info":[{"award-number":["ARS-47"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32273136"],"award-info":[{"award-number":["32273136"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["32273136"],"award-info":[{"award-number":["32273136"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"name":"BIO-KEY Germplasm Preservation Program","award":["HT-99982021-0392"],"award-info":[{"award-number":["HT-99982021-0392"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>Sperm activation and movement are essential in freshwater fishes, where limited energy reserves and a short motility duration require rapid ATP-driven motility regulated by osmolality and ions. However, the physiological and molecular mechanisms underlying osmolality and ion regulation in mandarin fish (Siniperca chuatsi) remain unclear, and ions in aquaculture systems may exert additional effects. Here, we integrated physiology with transcriptomic and proteomic analyses to characterize the effects of osmolality and ions on sperm motility and identify key genes, proteins, and regulatory networks involved in sperm activation and movement. Low osmolality (&lt;50 mOsm\/kg) promoted sperm activation, with sperm motility progressively decreasing as osmolality increased. Although relatively high osmolality supported motility maintenance, the initial motility was comparatively low, indicating the limited energy reserves of freshwater fish spermatozoa. Na+ and K+ activation media enhanced motility, whereas Ca2+ supplementation suppressed activation. Ca2+ levels below a 1:500 molar ratio (&lt;0.06 mM) did not significantly affect sperm motility, indicating the importance of limiting Ca2+ exposure during fertilization. Multi-omics analyses identified six differentially expressed genes, including grin3bb, cacna1c, and chrna7, and 35 differentially expressed proteins, including ATP1B, PRKCB, and CPT1A, as key molecules associated with Na+, K+, and Ca2+ signaling and energy metabolism. The calcium signaling pathway, cAMP signaling pathway, and oxidative phosphorylation were significantly enriched in KEGG enrichment analysis, while Gene Ontology (GO) enrichment analysis identified ion signaling and energy metabolism, including ion transport, the ATP metabolic process, and the glycolytic process. Our findings provide a putative regulatory network for sperm activation and movement in mandarin fish and a basis for optimizing artificial fertilization.<\/jats:p>","DOI":"10.3390\/ani16172762","type":"journal-article","created":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T09:20:43Z","timestamp":1788427243000},"page":"2762","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Integrating Physiology and Multi-Omics Reveals Mechanisms Underlying Sperm Activation and Movement in Mandarin Fish (Siniperca chuatsi)"],"prefix":"10.3390","volume":"16","author":[{"given":"Qinghua","family":"Wang","sequence":"first","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuxin","family":"Zhang","sequence":"additional","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhong","family":"Huang","sequence":"additional","affiliation":[{"name":"Shenzhen Base of South China Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, Shenzhen 518121, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weiwei","family":"Zhang","sequence":"additional","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yingxin","family":"Wu","sequence":"additional","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiajie","family":"Li","sequence":"additional","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yizheng","family":"Zhang","sequence":"additional","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lu","family":"Li","sequence":"additional","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhiming","family":"Zhu","sequence":"additional","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"},{"name":"China-ASEAN Belt and Road Joint Laboratory on Mariculture Technology, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5170-9613","authenticated-orcid":false,"given":"Zining","family":"Meng","sequence":"additional","affiliation":[{"name":"School of Life Sciences, State Key Laboratory of Biocontrol\/Guangdong Core Germplasm Bank for Marine Economic Animals, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)\/Guangdong Provincial Key Laboratory of Aquatic Economic Animals, Sun Yat-sen University, Guangzhou 510275, China"},{"name":"China-ASEAN Belt and Road Joint Laboratory on Mariculture Technology, Guangzhou 510275, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,2]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.cellbi.2005.06.004","article-title":"Sperm Motility in Fishes. (Ii) Effects of Ions and Osmolality: A Review","volume":"30","author":"Alavi","year":"2006","journal-title":"Cell Biol. Int."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1016\/j.theriogenology.2019.06.038","article-title":"Sperm Motility in Fishes: (iii) Diversity of Regulatory Signals from Membrane to the Axoneme","volume":"136","author":"Alavi","year":"2019","journal-title":"Theriogenology"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"362","DOI":"10.1111\/j.1439-0426.2008.01126.x","article-title":"Adaptation and Strategy for Fertilization in the Sperm of Teleost Fish","volume":"24","author":"Morisawa","year":"2008","journal-title":"J. Appl. Ichthyol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1016\/j.cellbi.2004.11.021","article-title":"Sperm Motility in Fishes. I. Effects of Temperature and Ph: A Review","volume":"29","author":"Alavi","year":"2005","journal-title":"Cell Biol. Int."},{"key":"ref_5","first-page":"726","article-title":"Sperm Motility of the Marine Teleosts Boops boops, Diplodus sargus, Mullus barbatus and Trachurus mediterraneus","volume":"52","author":"Lahnsteiner","year":"1998","journal-title":"J. Fish Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1016\/j.cbpa.2005.12.020","article-title":"Changes in Sperm Motility in Response to Osmolality\/Ca2+ in Three Indonesian Fresh Water Teleosts: Goby (Oxyeleotris marmorata), Java Carp (Puntius javanicus), and Catfish (Clarias batrachus)","volume":"143","author":"Morita","year":"2006","journal-title":"Comp. Biochem. Phys. A"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1175","DOI":"10.1242\/jcs.108.3.1175","article-title":"Change in Intracellular K+ Concentration Caused by External Osmolality Change Regulates Sperm Motility of Marine and Freshwater Teleosts","volume":"108","author":"Takai","year":"1995","journal-title":"J. Cell Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1054","DOI":"10.1016\/j.theriogenology.2008.11.006","article-title":"Influence of Osmolality and Ions on the Activation and Characteristics of Zebrafish Sperm Motility","volume":"71","author":"Kanuga","year":"2009","journal-title":"Theriogenology"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"630","DOI":"10.1242\/jeb.063628","article-title":"Transient Ca2+ Mobilization Caused by Osmotic Shock Initiates Salmonid Fish Sperm Motility","volume":"215","author":"Takei","year":"2012","journal-title":"J. Exp. Biol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"2052","DOI":"10.1073\/pnas.040558097","article-title":"Membrane Hyperpolarization Removes Inactivation of Ca2+ Channels, Leading to Ca2+ Influx and Subsequent Initiation of Sperm Motility in the Common Carp","volume":"97","author":"Krasznai","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1002\/jez.414","article-title":"Calcium-Induced Quiescence of Sperm Motility in the Bluegill (Lepomis macrochirus)","volume":"307A","author":"Zuccarelli","year":"2007","journal-title":"J. Exp. Zool. Part A"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/j.jprot.2018.08.020","article-title":"RNA-Seq Coupled to Proteomic Analysis Reveals High Sperm Proteome Variation between Two Closely Related Marine Mussel Species","volume":"192","author":"Romero","year":"2019","journal-title":"J. Proteom."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"111537","DOI":"10.1016\/j.isci.2024.111537","article-title":"Post-Testicular Spermatozoa of a Marine Teleost Can Conduct De Novo Cytoplasmic and Mitochondrial Translation","volume":"28","author":"Finn","year":"2025","journal-title":"iScience"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"e202400310","DOI":"10.1002\/pmic.202400310","article-title":"The Zebrafish Sperm Proteome","volume":"25","author":"Martins","year":"2025","journal-title":"Proteomics"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"14162","DOI":"10.1038\/s41598-022-18422-2","article-title":"Developmental Rna-Seq Transcriptomics of Haploid Germ Cells and Spermatozoa Uncovers Novel Pathways Associated with Teleost Spermiogenesis","volume":"12","author":"Dabad","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"147014","DOI":"10.1016\/j.gene.2022.147014","article-title":"Omics Insights into Spermatozoa Activation Induced by Fetal Bovine Serum in Viviparous Black Rockfish (Sebastes schlegelii)","volume":"851","author":"Niu","year":"2023","journal-title":"Gene"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1016\/j.theriogenology.2019.04.018","article-title":"Proteomic and Metabolomic Insights into the Functions of the Male Reproductive System in Fishes","volume":"132","author":"Dietrich","year":"2019","journal-title":"Theriogenology"},{"key":"ref_18","first-page":"10","article-title":"In-Depth Proteomic Analysis of Carp (Cyprinus carpio L.) Spermatozoa","volume":"12","author":"Dietrich","year":"2014","journal-title":"Comp. Biochem. Phys. D"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1569","DOI":"10.1002\/pmic.201300521","article-title":"Proteomic Identification of Rainbow Trout Sperm Proteins","volume":"14","author":"Nynca","year":"2014","journal-title":"Proteomics"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Chen, J., Zhou, A., Xie, S., Wang, C., Lv, Z., and Zou, J. (2016). Comparative Proteomic Identification of Mature and Immature Sperm in the Catfish Cranoglanis bouderius. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0151254"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.jprot.2017.01.019","article-title":"Proteomic Characterization of Seminal Plasma from Alternative Reproductive Tactics of Chinook Salmon (Oncorhynchus tswatchysha)","volume":"157","author":"Gombar","year":"2017","journal-title":"J. Proteom."},{"key":"ref_22","first-page":"113","article-title":"Classification and Geographical Distribution of Sinipercinae Fish in the Subfamily Sinipercae","volume":"2","author":"Zhou","year":"1988","journal-title":"Zool. Res."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1046\/j.1355-557x.2001.00006.x","article-title":"Weaning Chinese Perch Siniperca chuatsi (Basilewsky) onto Artificial Diets Based Upon Its Specific Sensory Modality in Feeding","volume":"32","author":"Liang","year":"2001","journal-title":"Aquac. Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"M1778","DOI":"10.1111\/1750-3841.12289","article-title":"Diversity of Lactic Acid Bacteria During Fermentation of a Traditional Chinese Fish Product, Chouguiyu (Stinky Mandarinfish)","volume":"78","author":"Dai","year":"2013","journal-title":"J. Food Sci."},{"key":"ref_25","unstructured":"Bureau of Fisheries, Ministry of Agriculture (2026). Chinese Fishery Statistical Yearbook. Chapter 2, Freshwater Aquaculture Production in Various Regions (by Species) (III), (In Chinese)."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"i884","DOI":"10.1093\/bioinformatics\/bty560","article-title":"Fastp: An Ultra-Fast All-in-One Fastq Preprocessor","volume":"34","author":"Chen","year":"2018","journal-title":"Bioinformatics"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1038\/nmeth.3317","article-title":"Hisat: A Fast Spliced Aligner with Low Memory Requirements","volume":"12","author":"Kim","year":"2015","journal-title":"Nat. Methods"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1038\/nbt.3122","article-title":"Stringtie Enables Improved Reconstruction of a Transcriptome from RNA-Seq Reads","volume":"33","author":"Pertea","year":"2015","journal-title":"Nat. Biotechnol."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Li, B., and Dewey, C.N. (2011). RSEM: Accurate Transcript Quantification from RNA-Seq Data with or without a Reference Genome. BMC Bioinform., 12.","DOI":"10.1186\/1471-2105-12-323"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated Estimation of Fold Change and Dispersion for RNA-Seq Data with Deseq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"D362","DOI":"10.1093\/nar\/gkw937","article-title":"The String Database in 2017: Quality-Controlled Protein\u2013Protein Association Networks, Made Broadly Accessible","volume":"45","author":"Szklarczyk","year":"2017","journal-title":"Nucleic Acids Res."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2498","DOI":"10.1101\/gr.1239303","article-title":"Cytoscape: A Software Environment for Integrated Models of Biomolecular Interaction Networks","volume":"13","author":"Shannon","year":"2003","journal-title":"Genome Res."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1126\/science.7444445","article-title":"Osmolality and Potassium Ion: Their Roles in Initiation of Sperm Motility in Teleosts","volume":"210","author":"Morisawa","year":"1980","journal-title":"Science"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1111\/j.1095-8649.2009.02504.x","article-title":"Frenetic Activation of Fish Spermatozoa Flagella Entails Short-Term Motility, Portending Their Precocious Decadence","volume":"76","author":"Cosson","year":"2010","journal-title":"J. Fish Biol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1023\/B:AQUI.0000017189.44263.bc","article-title":"The Ionic and Osmotic Factors Controlling Motility of Fish Spermatozoa","volume":"12","author":"Cosson","year":"2004","journal-title":"Aquacult. Int."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1242\/jeb.107.1.95","article-title":"Effects of Osmolality and Potassium on Motility of Spermatozoa from Freshwater Cyprinid Fishes","volume":"107","author":"Morisawa","year":"1983","journal-title":"J. Exp. Biol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1530\/jrf.0.1100315","article-title":"Effects of Extracellular Environment on the Osmotic Signal Transduction Involved in Activation of Motility of Carp Spermatozoa","volume":"110","author":"Gatti","year":"1997","journal-title":"J. Reprod. Fertil."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1111\/j.1439-0426.1996.tb00071.x","article-title":"Motility and Fertilizing Capacity of Fresh and Frozen-Thawed Spermatozoa in Sturgeons Acipenser Baeri and A. ruthenus","volume":"12","author":"Tsvetkova","year":"1996","journal-title":"J. Appl. Ichthyol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1238","DOI":"10.1111\/j.1365-2109.2004.01132.x","article-title":"Chemical Composition and Osmolality of Seminal Fluid of Acipenser Persicus; Their Physiological Relationship with Sperm Motility","volume":"35","author":"Alavi","year":"2004","journal-title":"Aquac. Res."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"492","DOI":"10.1111\/j.1095-8649.1995.tb01917.x","article-title":"Fine Structure and Motility of Spermatozoa and Composition of the Seminal Plasma in the Perch","volume":"47","author":"Lahnsteiner","year":"1995","journal-title":"J. Fish Biol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1111\/j.1439-0426.2008.01133.x","article-title":"Basic Physico-Chemical Parameters of Milt from Sea Trout (Salmo trutta M. trutta), Brook Trout (Salvelinus fontinalis) and Rainbow Trout (Oncorhynchus mykiss)","volume":"24","author":"Dziewulska","year":"2008","journal-title":"J. Appl. Ichthyol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"9325","DOI":"10.1073\/pnas.0611296104","article-title":"A Sperm-Specific Na+\/H+ Exchanger (Snhe) Is Critical for Expression and in Vivo Bicarbonate Regulation of the Soluble Adenylyl Cyclase (Sac)","volume":"104","author":"Wang","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"e07624","DOI":"10.7554\/eLife.07624","article-title":"A K+-Selective Cng Channel Orchestrates Ca2+ Signalling in Zebrafish Sperm","volume":"4","author":"Fechner","year":"2015","journal-title":"eLife"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"711","DOI":"10.21451\/1984-3143-AR2018-0024","article-title":"Na\/K-Atpase and Regulation of Sperm Function","volume":"15","author":"Thundathil","year":"2018","journal-title":"Anim. Reprod."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1093\/biolre\/ioaa093","article-title":"The Na+ and K+ Transport System of Sperm (Atp1a4) Is Essential for Male Fertility and an Attractive Target for Male Contraception","volume":"103","author":"Syeda","year":"2020","journal-title":"Biol. Reprod."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1083\/jcb.84.1.13","article-title":"Calcium-Induced Quiescence in Reactivated Sea Urchin Sperm","volume":"84","author":"Gibbons","year":"1980","journal-title":"J. Cell Biol."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Shoshan-Barmatz, V., Shteinfer-Kuzmine, A., and Verma, A. (2020). Vdac1 at the Intersection of Cell Metabolism, Apoptosis, and Diseases. Biomolecules, 10.","DOI":"10.3390\/biom10111485"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"9481","DOI":"10.1038\/s41467-025-65363-1","article-title":"Structural Basis of Apoptosis Induction by the Mitochondrial Voltage-Dependent Anion Channel","volume":"16","author":"Daniilidis","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"552","DOI":"10.1038\/nrm1150","article-title":"Regulation of Cell Death: The Calcium\u2013Apoptosis Link","volume":"4","author":"Orrenius","year":"2003","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"882","DOI":"10.1016\/j.bbadis.2015.01.019","article-title":"Calpain-1 Induces Endoplasmic Reticulum Stress in Promoting Cardiomyocyte Apoptosis Following Hypoxia\/Reoxygenation","volume":"1852","author":"Zheng","year":"2015","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"13783","DOI":"10.1073\/pnas.2001214117","article-title":"A Family of Hyperpolarization-Activated Channels Selective for Protons","volume":"117","author":"Wobig","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Soda, T., Brunetti, V., Berra-Romani, R., and Moccia, F. (2023). The Emerging Role of N-Methyl-D-Aspartate (Nmda) Receptors in the Cardiovascular System: Physiological Implications, Pathological Consequences, and Therapeutic Perspectives. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24043914"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"22754","DOI":"10.1074\/jbc.271.37.22754","article-title":"Identification of the Mammalian Na,K-Atpase \u03923 Subunit","volume":"271","author":"Malik","year":"1996","journal-title":"J. Biol. Chem."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"17024","DOI":"10.1073\/pnas.0606539103","article-title":"Atherosclerosis-Related Molecular Alteration of the Human Cav1.2 Calcium Channel A1c Subunit","volume":"103","author":"Tiwari","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"199","DOI":"10.2147\/OAJC.S572080","article-title":"Calcium Channels and Modulators as Potential Therapeutic Targets for Contraceptives and Male Fertility: A Scoping Review","volume":"16","author":"Twambaze","year":"2025","journal-title":"Open Access J. Contracept."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"807","DOI":"10.1095\/biolreprod.105.042184","article-title":"Mice Deficient in Chrna7, a Subunit of the Nicotinic Acetylcholine Receptor, Produce Sperm with Impaired Motility","volume":"73","author":"Bray","year":"2005","journal-title":"Biol. Reprod."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1016\/j.neuropharm.2015.02.006","article-title":"The Human Chrna7 and Chrfam7a Genes: A Review of the Genetics, Regulation, and Function","volume":"96","author":"Sinkus","year":"2015","journal-title":"Neuropharmacology"},{"key":"ref_58","first-page":"E395","article-title":"Protein Kinase C: Poised to Signal","volume":"298","author":"Newton","year":"2009","journal-title":"Am. J. Physiol.-Endoc. M."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1017\/S0967199400003932","article-title":"Inositol Triphosphate Receptors in Sea Urchin Sperm","volume":"5","author":"Zapata","year":"1997","journal-title":"Zygote"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"605","DOI":"10.1038\/nature00944","article-title":"Structural Changes in the Calcium Pump Accompanying the Dissociation of Calcium","volume":"418","author":"Toyoshima","year":"2002","journal-title":"Nature"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"33742","DOI":"10.1074\/jbc.M404628200","article-title":"Targeted Ablation of Plasma Membrane Ca2+-Atpase (Pmca) 1 and 4 Indicates a Major Housekeeping Function for Pmca1 and a Critical Role in Hyperactivated Sperm Motility and Male Fertility for Pmca4","volume":"279","author":"Okunade","year":"2004","journal-title":"J. Biol. Chem."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1016\/j.ejcb.2007.02.003","article-title":"Localization and Regulation of Plasma Membrane Ca2+-Atpase in Bovine Spermatozoa","volume":"86","author":"Triphan","year":"2007","journal-title":"Eur. J. Cell Biol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"14681","DOI":"10.1073\/pnas.95.25.14681","article-title":"Bax Interacts with the Permeability Transition Pore to Induce Permeability Transition and Cytochrome C Release in Isolated Mitochondria","volume":"95","author":"Narita","year":"1998","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/j.repbio.2013.12.005","article-title":"Motility of Fish Spermatozoa: From External Signaling to Flagella Response","volume":"14","author":"Dzyuba","year":"2014","journal-title":"Reprod. Biol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1016\/j.aquaculture.2016.05.038","article-title":"Energetics of Fish Spermatozoa: The Proven and the Possible","volume":"472","author":"Dzyuba","year":"2017","journal-title":"Aquaculture"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"1615","DOI":"10.1007\/s10695-019-00650-5","article-title":"Bioenergetics of Fish Spermatozoa with Focus on Some Herring (Clupea harengus) Enzymes","volume":"45","author":"Gronczewska","year":"2019","journal-title":"Fish Physiol. Biochem."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"741538","DOI":"10.1016\/j.aquaculture.2024.741538","article-title":"Integrated Sperm Movement, Proteomics, and Phosphoproteomics Elucidate the Roles of Osmolality, Ions, and Key Proteins in Sperm Activation of Nile Tilapia (Oreochromis niloticus)","volume":"595","author":"Wang","year":"2025","journal-title":"Aquaculture"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"454","DOI":"10.1002\/(SICI)1097-010X(19990901)284:4<454::AID-JEZ12>3.0.CO;2-O","article-title":"Sperm Metabolism of the Telost Fishes Chalcalburnus chalcoides and Oncorhynchus mykiss and Its Relation to Motility and Viability","volume":"284","author":"Lahnsteiner","year":"1999","journal-title":"J. Exp. Zool."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1016\/j.jprot.2016.02.029","article-title":"Motility of Carp Spermatozoa Is Associated with Profound Changes in the Sperm Proteome","volume":"138","author":"Dietrich","year":"2016","journal-title":"J. Proteom."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1007\/s10695-024-01418-2","article-title":"Evaluation of Carp Sperm Respiration: Fluorometry with Optochemical Oxygen Sensor Versus Polarography","volume":"51","author":"Musatova","year":"2024","journal-title":"Fish Physiol. Biochem."}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/16\/17\/2762\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T09:36:59Z","timestamp":1788428219000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/16\/17\/2762"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,2]]},"references-count":70,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["ani16172762"],"URL":"https:\/\/doi.org\/10.3390\/ani16172762","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,2]]}}}