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Spermatic mitochondria: role in oxidative homeostasis, sperm function and possible tools for their assessment

  • João Diego de Agostini Losano (a1), Daniel de Souza Ramos Angrimani (a1), Roberta Ferreira Leite (a1), Bárbara do Carmo Simões da Silva (a1), Valquíria Hyppolito Barnabe (a1) and Marcilio Nichi (a1)...


Despite sperm mitochondrial relevance to the fertilization capacity, the processes involved in the production of ATP and functional dynamics of sperm mitochondria are not fully understood. One of these processes is the paradox involved between function and formation of reactive oxygen species performed by the organelle. Therefore, this review aimed to provide data on the role of sperm mitochondria in oxidative homeostasis and functionality as well the tools to assess sperm mitochondrial function.


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Author for correspondence: Department of Animal Reproduction, College of Veterinary Medicine and Animal Science, University of São Paulo, Av. Prof. Orlando Marques de Paiva, 87-05508-270, São Paulo, Brazil. Tel: +55 11 30911423. Fax: +55 11 30911437. E-mail:


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Agarwal, A, Nallella, KP, Allamaneni, SSR Said, TM (2004) Role of antioxidants in treatment of male infertility: an overview of the literature. Reprod BioMed Online 8, 616627.
Agarwal, A, Virk, G, Ong, C du Plessis, SS (2014) Effect of oxidative stress on male reproduction. World J Mens Health 32, 117.
Aitken, RJ (2006) Sperm function tests and fertility. Int J Androl 29, 6975.
Aitken, RJ, Paterson, M, Fisher, H, Buckingham, DW van Duin, M (1995) Redox regulation of tyrosine phosphorylation in human spermatozoa and its role in the control of human sperm function. J Cell Sci 108, 20172025.
Aitken, RJ, Ryan, AL, Baker, MA McLaughlin, EA (2004) Redox activity associated with the maturation and capacitation of mammalian spermatozoa. Free Rad Biol Med 36, 9941010.
Alvarez, JG, Touchstone, JC, Blasco, L Storey, BT (1987) Spontaneous lipid peroxidation and production of hydrogen peroxide and superoxide in human spermatozoa superoxide dismutase as major enzyme protectant against oxygen toxicity. J Androl 8, 338348.
Amaral, A, Lourenço, B, Marques, M Ramalho-Santos, J (2013) Mitochondria functionality and sperm quality. Reproduction 146, R163R174.
Askari, HA, Check, JH, Peymer, N Bollendorf, A (1994) Effect of natural antioxidants tocopherol and ascorbic acids in maintenance of sperm activity during freeze-thaw process. Syst Biol Reprod Med 33, 1115.
Bailey, S Macardle, PJ (2006) A flow cytometric comparison of indo-1 to fluo-3 and Fura Red excited with low power lasers for detecting Ca2+ flux. J Immunol Methods 311, 220225.
Barros, P (2007) Estresse oxidativo e integridade do DNA em sêmen resfriado de gato-do-mato-pequeno (Leopardus tigrinus SCHREBER 1775).
Bilodeau, J.-F, Chatterjee, S, Sirard, M.-A Gagnon, C (2000) Levels of antioxidant defenses are decreased in bovine spermatozoa after a cycle of freezing and thawing. Mol Reprod Dev 55, 282288.
Bilodeau, JF, Blanchette, S, Gagnon, C Sirard, MA (2001) Thiols prevent H2O2-mediated loss of sperm motility in cryopreserved bull semen. Theriogenology 56, 275286.
Blumer, CG, Restelli, AE, Giudice, P.T.D, Soler, TB, Fraietta, R, Nichi, M, Bertolla, RP Cedenho, AP (2012) Effect of varicocele on sperm function and semen oxidative stress. BJU Int 109, 259265.
Brand, MD Esteves, TC (2005) Physiological functions of the mitochondrial uncoupling proteins, UCP2 and, UCP3. Cell Metab 2, 8593.
Breitbart, H (2002) Intracellular calcium regulation in sperm capacitation and acrosomal reaction. Mol Cell Endocrinol 187, 139144.
Brewis, IA, Morton, IE, Mohammad, SN, Browes, CE Moore, H.D.M (2000) Measurement of intracellular calcium concentration and plasma membrane potential in human spermatozoa using flow cytometry. J Androl 21, 238249.
Celeghini, E.C.C, De Arruda, RP, De Andrade, A.F.C, Nascimento, J Raphael, CF (2007) Practical techniques for bovine sperm simultaneous fluorimetric assessment of plasma., acrosomal and mitochondrial membranes. Reprod Domest Anim 42, 479488.
Chazotte, B (2011) Labeling mitochondria with MitoTracker dyes. Cold Spring Harb Protoc 8, 990992.
Chen, LB (1988) Mitochondrial membrane potential in living cells. Ann Rev Cell Biol 4, 155181.
Christen, R, Schackmann, RW Shapiro, BM (1983) Metabolism of sea urchin sperm., Interrelationships between intracellular pH, ATPase activity and mitochondrial respiration. J Biol Chem 258, 53925399.
Collin, S, Sirard, MA, Dufour, M Bailey, JL (2000) Sperm calcium levels and chlortetracycline fluorescence patterns are related to the in vivo fertility of cryopreserved bovine semen. J Androl. 21, 938943.
Copeland, WC (2002) Mitochondrial DNA, Springer.
Cummins, J (1998) Mitochondrial, DNA in mammalian reproduction. Rev Reprod 3, 172182.
de Lamirande, E Cagnon, C (1993) Human sperm hyperactivation and capacitation as parts of an oxidative process. Free Rad Biol Med 14, 157166.
de Lamirande, E, Jiang, H, Zini, A, Kodama, H Gagnon, C (1997) Reactive oxygen species and sperm physiology. Rev Reprod 2, 4854.
de Lamirande, E.V.E, Tsai, C, Harakat, A Gagnon, C (1998) Involvement of reactive oxygen species in human sperm acrosome reaction induced by A23187, lysophosphatidylcholine, and biological fluid ultrafiltrates. J Androl 19, 585594.
Del Olmo, E, Bisbal, A, Maroto-Morales, A, García-Alvarez, O, Ramon, M, Jimenez-Rabadan, P, Martínez-Pastor, F, Soler, AJ, Garde, JJ Fernández-santos, MR (2013) Fertility of cryopreserved ovine semen is determined by sperm velocity. Anim Reprod Sci 138, 102109.
Dorweiler, B, Pruefer, D, Andrasi, T, Maksan, S, Schmiedt, W, Neufang, A Vahl, C (2007) Ischemia-reperfusion injury. Eur J Trauma Emerg Surg 33, 600612.
Fernández-Santos, MR, Martínez-Pastor, F, García-Macías, V, Esteso, MC, Soler, AJ, Paz, P, Anel, L Garde, JJ (2007) Sperm characteristics and, D.NA integrity of, Iberian red deer (Cervus elaphus hispanicus) epididymal spermatozoa frozen in the presence of enzymatic and nonenzymatic antioxidants. J Androl 28, 294305.
Ford, W.C.L (2006) Glycolysis and sperm motility: does a spoonful of sugar help the flagellum go round? Hum Reprod Update 12, 269274.
Forster, S, Thumser, AE, Hood, SR Plant, N (2012) Characterization of rhodamine-123 as a tracer dye for use in in vitro drug transport assays. PLoS One 7, e33253.
Gallon, F, Marchetti, C, Jouy, N Marchetti, P (2006) The functionality of mitochondria differentiates human spermatozoa with high and low fertilizing capability. Fertil Steril 86, 15261530.
García-Rivas, G, de, J, Carvajal, K, Correa, F Zazueta, C (2006) Ru360, a specific mitochondrial calcium uptake inhibitor., improves cardiac post-ischaemic functional recovery in rats in vivo . Brit J Pharmacol 149, 829837.
Garner, DL, Thomas, CA, Joerg, HW, DeJarnette, JM Marshall, CE (1997) Fluorometric assessments of mitochondrial function and viability in cryopreserved bovine spermatozoa. Biol Reprod 57, 14011406.
Gillan, L, Evans, G Maxwell, WMC (2005) Flow cytometric evaluation of sperm parameters in relation to fertility potential. Theriogenology 63, 445457.
Giojalas, LC (1998) Correlation between response to progesterone and other functional parameters in human spermatozoa. Fertil Steril 69, 107111.
Graham, JK, Kunze, E Hammerstedt, RH (1990) Analysis of sperm cell viability., acrosomal integrity, and mitochondrial function using flow cytometry. Biol Reprod 43, 5564.
Gunter, TE, Yule, DI, Gunter, KK, Eliseev, RA Salter, JD (2004) Calcium and mitochondria. FEBS Lett 567, 96102 2004.
Hallap, T, Nagy, S, Jaakma, U, Johannisson, A Rodriguez-Martinez, H (2005) Mitochondrial activity of frozen–thawed spermatozoa assessed by MitoTracker Deep Red 633. Theriogenology 63, 23112322.
Halliwell, B (1999) Free Radicals in Biology and Medicine, Oxford University Press.
Halliwell, B Gutteridge, J (1985) Free Radicals in Biology and Medicine, Pergamon.
Hammerstedt, RH, Graham, JK Nolan, JP (1990) cryopreservation of mammalian sperm: what we ask them to survive. J Androl 11, 7388.
Harrison, R.A.P, Mairet, B Miller, N.G.A (1993) Flow cytometric studies of bicarbonate-mediated, Ca2+ influx in boar sperm populations. Mol Reprod Dev 35, 197208.
Herzog, V Fahimi, HD (1973) A new sensitive colorimetric assay for peroxidase using 3,3′-diaminobenzidine as hydrogen donor. Anal Biochem 55, 554562.
Holt, WV (2000) Fundamental aspects of sperm cryobiology: the importance of species and individual differences. Theriogenology 53, 4758.
Hrudka, F (1987) Cytochemical and ultracytochemical demonstration of cytochrome c oxidase in spermatozoa and dynamics of its changes accompanying ageing or induced by stress. Int J Androl 10, 809828.
Hu, CH, Zhuang, XJ, Wei, YM, Zhang, M, Lu, SS, Lu, YG, Yang, XG Lu, KH (2017) comparison of mitochondrial function in boar and bull spermatozoa throughout cryopreservation based on JC-1 staining. Cryo Lett 38, 7579.
Irvine, DS Aitken, RJ (1986) Measurement of intracellular calcium in human spermatozoa. Gamete Res 15, 5771.
John, J.C.S, Jokhi, RP Barratt, C.L.R (2005) The effect of mitochondrial genetics on male infertility. Int J Androl 28, 6573.
Kasai, T, Ogawa, K, Mizuno, K, Nagai, S, Uchida, Y, Ohta, S, Fujie, M, Suzuki, K, Hirata, S Hoshi, K (2002) Relationship between sperm mitochondrial membrane potential, sperm motility, and fertility potential. Asian J Androl 4, 97104.
Kasianowicz, J, Benz, R McLaughlin, S (1984) The kinetic mechanism by which CCCP (carbonyl cyanidem-chlorophenylhydrazone) transports protons across membranes. J Membr Biol 82, 179190.
Koppers, AJ, De Iuliis, GN, Finnie, JM, McLaughlin, EA Aitken, RJ (2008) Significance of mitochondrial reactive oxygen species in the generation of oxidative stress in spermatozoa. J Clin Endocrinol Metab 93, 31993207.
Krzyzosiak, J, Molan, P Vishwanath, R (1999) Measurements of bovine sperm velocities under true anaerobic and aerobic conditions. Anim Reprod Sci 55, 163173.
Lardy, H, Winchester, B Phillips, P (1945) The respiratory metabolism of ram spermatozoa. Arch Biochem 6, 3340.
Lin, MT Beal, MF (2006) Mitochondrial dysfunction and oxidative stress in neurodegenerative diseases. Nature 443, 787795.
Losano, JD, Padín, JF, Méndez-López, I, Angrimani, DS, García, AG, Barnabe, VH et al. 2017a) The stimulated glycolytic pathway is able to maintain ATP levels and kinetic patterns of bovine epididymal sperm subjected to mitochondrial uncoupling. oxidative medicine and cellular longevity. Oxid Med Cell Longev 2017, 1682393.
Losano, J, Angrimani, D, Dalmazzo, A, Rui, B, Brito, M, Mendes, C et al. 2017b) Effect of mitochondrial uncoupling and glycolysis inhibition on ram sperm functionality. Reprod Domest Anim 52, 289297.
Lowell, BB Shulman, GI (2005) Mitochondrial dysfunction and type 2 diabetes. Science 307, 384397.
Machado-Oliveira, G, Lefièvre, L, Ford, C, Herrero, MB, Barratt, C, Connolly, TJ, Nash, K, Morales-García, A, Kirkman-Brown, J Publicover, S (2008) Mobilisation of Ca2+ stores and flagellar regulation in human sperm by S-nitrosylation: a role for NO synthesised in the female reproductive tract. Development 135, 36773686.
Mahanes, MS, Ochs, DL Eng, LA (1986) Cell calcium of ejaculated rabbit spermatozoa before and following in vitro capacitation. Biochem Biophys Res Commun 134, 664670.
Mailloux, RJ Harper, M.-E (2011) Uncoupling proteins and the control of mitochondrial reactive oxygen species production. Free Rad Biol Med 51, 11061115.
Marchetti, C, Jouy, N, Leroy-Martin, B, Defossez, A, Formstecher, P Marchetti, P (2004) Comparison of four fluorochromes for the detection of the inner mitochondrial membrane potential in human spermatozoa and their correlation with sperm motility. Hum Reprod 19, 22672276.
Marchetti, C, Obert, G, Deffosez, A, Formstecher, P Marchetti, P (2002) Study of mitochondrial membrane potential reactive oxygen species, DNA fragmentation and cell viability by flow cytometry in human sperm. Hum Reprod 17, 12571265.
Margulis, L (1970) Origin of Eukaryotic Cells. Evidence and Research Implications for a Theory of the Origin and Evolution of Microbial, Plant, and Animal Cells on the Precambrian Earth. Yale, University, Press: New Haven.
McCormack, JG Denton, RM (1993) Mitochondrial Ca2+ transport and the role of intramitochondrial Ca2+ in the regulation of energy metabolism. Dev Neurosci 15, 165173.
McCormack, JG, Halestrap, AP Denton, RM (1990) Role of calcium ions in regulation of mammalian intramitochondrial metabolism. Physiol Rev 70, 391425.
Meijer, J Fentener, Van Vlissingen, J (1993) Gross structure and development of reproductive organs. In Reproduction in Domesticated Animals, (ed. G.L. King), pp. 9–26. World Animal, Science-B9. Amsterdam: Elsevier Science Publishers.
Merritt, JE, McCarthy, SA, Davies, MP Moores, KE (1990) Use of fluo-3 to measure cytosolic Ca2+ in platelets and neutrophils, loading cells with the dye, calibration of traces., measurements in the presence of plasma, and buffering of cytosolic Ca2+ . Biochem J 269, 513519.
Modica-Napolitano, JS Singh, KK (2004) Mitochondrial dysfunction in cancer. Mitochondrion 4, 755762.
Mukai, C Okuno, M (2004) glycolysis plays a major role for adenosine triphosphate supplementation in mouse sperm flagellar movement. Biol Reprod 71, 540547.
Nascimento, JM, Shi, LZ, Tam, J, Chandsawangbhuwana, C, Durrant, B, Botvinick, EL Berns, MW (2008) Comparison of glycolysis and oxidative phosphorylation as energy sources for mammalian sperm motility, using the combination of fluorescence imaging., laser tweezers., and real-time automated tracking and trapping. J Cell Physiol 217, 745751.
Nelson, DL Cox, MM (2008) Principles of Biochemistry, 5th edn, New York, NY: W.H. Freeman and Company.
Nevo, AC Rikmenspoel, R (1970) Diffusion of ATP in sperm flagella. J Theoret Biol 26, 1118.
Nichi, M, Bols, P.E.J, Züge, RM, Barnabe, VH, Goovaerts, I.G.F, Barnabe, RC Cortada, C.N.M (2006) Seasonal variation in semen quality in Bos indicus and Bos taurus bulls raised under tropical conditions. Theriogenology 66, 822888.
Nichi, M, Goovaerts, IG, Cortada, CN, Barnabe, VH, De Clercq, JB Bols, PE (2007a) Roles of lipid peroxidation and cytoplasmic droplets on in vitro fertilization capacity of sperm collected from bovine epididymides stored at 4 and 34 degrees C. Theriogenology 67, 334340.
Nichi, M, Goovaerts, IGF, Cortada, C.N.M, Barnabe, VH, De Clercq, J.B.P Bols, P.E.J (2007b) Roles of lipid peroxidation and cytoplasmic droplets on in vitro fertilization capacity of sperm collected from bovine epididymides stored at 4 and 34°C. Theriogenology 67, 334340.
Nordberg, J Arnér, E.S.J (2001) Reactive oxygen species., antioxidants., and the mammalian thioredoxin system. Free Rad Biol Med 31, 12871312.
O’Connell, M, McClure, N Lewis, S.E.M (2002) The effects of cryopreservation on sperm morphology., motility and mitochondrial function. Hum Reprod 17, 704709.
Paul, C, Teng, S Saunders, P.T.K (2009) A single, mild, transient scrotal heat stress causes hypoxia and oxidative stress in mouse testes, which induces germ cell death. Biol Reprod 80, 913919.
Pariz, JR Hallak, J (2016) Effects of caffeine supplementation in post-thaw human semen over different incubation periods. Andrologia 48, 961966.
Penefsky, HS (1985) Mechanism of inhibition of mitochondrial adenosine triphosphatase by dicyclohexylcarbodiimide and oligomycin: relationship to ATP synthesis. Proc Natl Acad Sci USA 82, 15891593.
Perchec, G, Jeulin, C, Cosson, J, Andre, F Billard, R (1995) Relationship between sperm ATP content and motility of carp spermatozoa. J Cell Sci 108, 747753.
Piccoli, C, Scrima, R, D’Aprile, A, Ripoli, M, Lecce, L, Boffoli, D Capitanio, N (2006) Mitochondrial dysfunction in hepatitis C virus infection. Biochim Biophys Acta 1757, 14291437.
Poot, M, Zhang, YZ, Krämer, JA, Wells, KS, Jones, LJ, Hanzel, DK, Lugade, AG, Singer, VL Haugland, RP (1996) Analysis of mitochondrial morphology and function with novel fixable fluorescent stains. J Histochem Cytochem 44, 13631372.
Ramalho-Santos, J, Varum, S, Amaral, S, Mota, PC, Sousa, AP Amaral, A (2009) Mitochondrial functionality in reproduction: from gonads and gametes to embryos and embryonic stem cells. Hum Reprod Update 15, 553572.
Rasola, A Geuna, M (2001) A flow cytometry assay simultaneously detects independent apoptotic parameters. Cytometry 45, 151157.
Reyes, JG, Farias, JG, Henríquez-Olavarrieta, S, Madrid, E, Parraga, M, Zepeda, AB Moreno, RD (2012) The hypoxic testicle: physiology and pathophysiology. Oxid Med Cell Longev 2012, 929285.
Rowe, M, Laskemoen, T, Johnsen, A Lifjeld, JT (2013) Evolution of sperm structure and energetics in passerine birds. Proc Biol Sci 280 (1753):20122616.
Ruiz-Pesini, E, Lapeña, A.-C, Díez-Sánchez, C, Pérez-Martos, A, Montoya, J, Alvarez, E, Díaz, M, Urriés, A, Montoro, L, López-Pérez, MJ Enríquez, JA (2000) Human mtDNA haplogroups associated with high or reduced spermatozoa motility. Am J Hum Genet 67, 682696.
Samizo, K, Ishikawa, R, Nakamura, A Kohama, K (2001) A highly sensitive method for measurement of myosin atpase activity by reversed-phase high-performance liquid chromatography. Anal Biochem 293, 212215.
Saraste, M (1999) Oxidative phosphorylation at the fin de siècle. Science 283, 14881493.
Sariozkan, S, Bucak, MN, Tuncer, PB, Ulutas, PA Bilgen, A (2009) The influence of cysteine and taurine on microscopic-oxidative stress parameters and fertilizing ability of bull semen following cryopreservation. Cryobiology 58, 134138.
Schober, D, Aurich, C, Nohl, H Gille, L (2007) Influence of cryopreservation on mitochondrial functions in equine spermatozoa. Theriogenology 68, 745754.
Sherer, TB, Betarbet, R, Testa, CM, Seo, BB, Richardson, JR, Kim, JH, Miller, GW, Yagi, T, Matsuno-Yagi, A Greenamyre, JT (2003) Mechanism of toxicity in rotenone models of Parkinson’s disease. J Neurosci 23, 1075610764.
Simões, R, Feitosa, WB, Siqueira, A.F.P, Nichi, M, Paula-Lopes, FF, Marques, MG, Peres, MA, Barnabe, VH, Visintin, JA Assumpção, M.E.O (2013) Influence of bovine sperm, DNA fragmentation and oxidative stress on early embryo in vitro development outcome. Reproduction 146, 433441.
Slater, EC (1973) The mechanism of action of the respiratory inhibitor, antimycin. Biochim Biophys Acta 301, 129154.
Smiley, ST, Reers, M, Mottola-Hartshorn, C, Lin, M, Chen, A, Smith, TW, Steele, GD Chen, LB (1991) Intracellular heterogeneity in mitochondrial membrane potentials revealed by a, J.-aggregate-forming lipophilic cation, J.C-1. Proc Natl Acad Sci USA 88, 36713675.
St John, J, Bowles, EJ Amaral, A (2006) Sperm mitochondria and fertilisation. Soc Reprod Fertil Suppl 65, 399416.
St John, JC (2002) The transmission of mitochondrial, DNA following assisted reproductive techniques. Theriogenology 57, 109123.
Storey, BT (2008) Mammalian sperm metabolism: oxygen and sugar, friend and foe. Int J Dev Biol 52, 427.
Szalai, G, Krishnamurthy, R Hajnóczky, G (1999) Apoptosis driven by IP3-linked mitochondrial calcium signals. EMBO J 18, 63496361.
Taylor, K, Roberts, P, Sanders, K Burton, P (2009) Effect of antioxidant supplementation of cryopreservation medium on post-thaw integrity of human spermatozoa. Reprod BioMed Online 18, 184189.
Terada, H (1990) Uncouplers of oxidative phosphorylation. Environ Health Perspect 87, 213.
Territo, PR, French, SA, Dunleavy, MC, Evans, FJ Balaban, RS (2001) Calcium activation of heart mitochondrial oxidative phosphorylation: rapid kinetics of mv, O2, NADH, and light scattering. J Biol Chem 276, 25862599.
Thomson, LK, Fleming, SD, Aitken, RJ, De Iuliis, GN, Zieschang, JA Clark, AM (2009) Cryopreservation-induced human sperm, DNA damage is predominantly mediated by oxidative stress rather than apoptosis. Hum Reprod 24, 20612070.
Thu, LT, Ahn, JR Woo, S.-H (2006) Inhibition of L-type Ca2+ channel by mitochondrial, Na+–Ca2+ exchange inhibitor CGP-37157 in rat atrial myocytes. Eur J Pharmacol 552, 1519.
Travis, AJ, Foster, JA, Rosenbaum, NA, Visconti, PE, Gerton, GL, Kopf, GS Moss, SB (1998) Targeting of a germ cell-specific type 1 hexokinase lacking a porin-binding domain to the mitochondria as well as to the head and fibrous sheath of murine spermatozoa. Mol Biol Cell 9, 263276.
Troiano, L, Granata, ARM, Cossarizza, A, Kalashnikova, G, Bianchi, R, Pini, G, Tropea, F, Carani, C Franceschi, C (1998) mitochondrial membrane potential and DNA stainability in human sperm cells: a flow cytometry analysis with implications for male infertility. Exp Cell Res 241, 384393.
Turner, RM (2003) Tales from the tail: what do we really know about sperm motility? J Androl 24, 790803.
Vernet, P, Aitken, RJ Drevet, JR (2004) Antioxidant strategies in the epididymis. Mol Cell Endocrinol 216, 3139.
Vincent, AM, Olzmann, JA, Brownlee, M, Sivitz, WI Russell, JW (2004) Uncoupling proteins prevent glucose-induced neuronal oxidative stress and programmed cell death. Diabetes 53, 726734.
Wang, G, Guo, Y, Zhou, T, Shi, X, Yu, J, Yang, Y, Wu, Y, Wang, J, Liu, M, Chen, X, Tu, W, Zeng, Y, Jiang, M, Li, S, Zhang, P, Zhou, Q, Zheng, B, Yu, C, Zhou, Z, Guo, X Sha, J (2013) In-depth proteomic analysis of the human sperm reveals complex protein compositions. J Proteomics 79, 114122.
Wang, X, Sharma, RK, Gupta, A, George, V, Thomas, AJ Jr, Falcone, T Agarwal, A (2003) Alterations in mitochondria membrane potential and oxidative stress in infertile men: a prospective observational study. Fertil Steril 80 (Suppl 2), :844850.
White, IG Wales, RG (1961) Comparison of epididymal and ejaculated semen of the ram. J Reprod Fertil 2, 225237.
Wojcik, C, Sawicki, W, Marianowski, P, Benchaib, M, Czyba, JC Guerin, JF (2000) Cyclodextrin enhances spermicidal effects of magainin-2-amide. Contraception 62, 99103.
Zhang, CL Wu, BJ (1996) Development of calcium fluorescent probes and their application in life sciences. Progr Physiol 27, 3742.
Zapzalka, DM, Redmon, JB Pryor, JL (1999) A survey of oncologists regarding sperm cryopreservation and assisted reproductive techniques for male cancer patients. Cancer 86, 18121817.



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