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Selenium/Selenoproteins in Male Reproductive Function
Selenium (Se) is an important trace mineral having many essential roles at the cellular and organismal levels in animal and human health. The biological effects of Se are mainly carried out by selenoproteins (encoded by 25 genes in humans and 24 in mice). As an essential component of selenoproteins, Se performs structural and enzymic roles; in the latter context it is well known for its catalytic and antioxidative functions. Studies involving different animal models have added great value to our understanding regarding the potential implications of Se and selenoproteins in mammalian fertility and reproduction.
1.2. Implication of Se in Mammalian Reproduction: An Overview
2. Important Selenoproteins Relevant to Male Reproduction
|Selenoprotein Gene *||Symbol ||General Description/Function
|mRNA *||Protein *||Relevance to Male Reproductive Function|
|Glutathione peroxidase 4||Gpx4||Detoxification of lipid hydroperoxides, Antioxidant in membranes, functions as structural protein in sperm, also implicated in apoptosis||++++||++++||Structural protein of sperm midpiece mitochondrial sheath and involved in sperm chromatin condensation . Implication in male fertility .|
|Thioredoxin-glutathione reductase||Txnrd3 (TGR)||Part of the thioredoxin system, Antioxidant function, redox regulation, cell signaling||+||+||Implicated in formation of disulfide bond and sperm maturation process . Expressed in post-pubertal testis, particularly abundant in elongated spermatids at the site of mitochondrial sheath formation .|
|Selenoprotein P||Selenop||Primarily responsible for Se transport and also performs antioxidative role.
Considered as a major contributor to plasma Se and a reliable biomarker of Se status. Its deficiency causes infertility characterized by abnormal sperm in mice
|+||+||Implicated in male fertility .
Implicated in transport of Se to spermatogenic cells .
Essential for sperm development in mice .
|Selenoprotein V||Selenov||Largely unidentified, potential role in redox regulation||+||n.d.||Specifically expressed in rodent testes . In situ hybridization trials have demonstrated the expression of Selenov mRNA in seminiferous tubules in mouse, however, its precise function in spermatogenesis is largely unexplored .|
|Selenoprotein W||Selenow||Antioxidant protection||+||+||n.d. *|
|Selenoprotein K||Selenok||Possible antioxidant protection in cardiomyocytes, Endoplasmic reticulum transmembrane protein||++||n.d.||n.d. *|
|Selenoprotein F||Selenof||Role in cell apoptosis and mediation of chemo-preventive effects of Se||+||n.d.||n.d. *|
|Selenoprotein S||Selenos||Cellular redox balance,
Possible influence in inflammatory response
|Selenophosphate synthetase 2||Sephs2||Required for biosynthesis of selenophosphate, a precursor of selenocysteine, and thus for selenoprotein synthesis||+||n.d.||n.d.|
3. Role of Selenium in Male Reproduction
3.1. Role of Selenium in Steroidogenesis and Spermatogenesis
3.2. Implication of Se on Male Fertility-Related Parameters
|Model||Treatment||Key Observations Reported||Ref.|
|Sprague-Dawley rats||Se nanoparticles at supranutritional levels (0.2, 0.4, or 0.8 mg Se per kg body weight)||Sperm parameters such as, sperm concentration, motility, and morphological features were all improved at supranutritional levels. However, these parameters were significantly affected when rats were supplemented with higher levels (nonlethal level) of Se nanoparticles i.e., at 2.0, 4.0, or 8.0 mg Se per kg body weight.|||
|Sprague-Dawley rats||Treated with inorganic Se [0.01(deficient); 0.25 (adequate); 3 (excess); or 5 (excess) mg per kg] for four weeks||The U-shaped response of dietary Se was observed on DNA damage and sperm quality. Se deficiency showed a lower expression of sensitive antioxidant selenoproteins (Gpx1 and Txnrds). However, excessive doses of Se impaired sperm quality and this was linked with reduced mRNA expression of nGpx4.|||
|Mouse||Se-supplement (inorganic Se (0.3 μg/g Se) or organic Se-enriched probiotics (containing 0.3 μg/g Se) given for 75 days||Organic Se co-supplemented with probiotics significantly improved male fertility in mice. The ameliorated fertility index included the parameters such as, reduced testicular tissue injury, increased levels of serum testosterone, and improved sperm indices in Se-supplemented group. As such, these improved fertility-related parameters were ascribed to be the result of the antioxidant function of Se.|||
|Mouse||0.2 ppm sodium selenite;
1.0 ppm sodium selenite
|Mice in both groups showed an increased occurrence of mitochondria- and plasma membrane-related defects, and DNA damage in sperm. However, these damages were more pronounced in mice exposed to Se-deficient feed.|||
|Mouse||Se-deficient diet (0.02 ppm)
Se-sufficient (0.2 ppm); organic Se
|Sperm from Se-deficient mice demonstrated vitiated chromatin condensation, declined in vitro fertilization ability and increased lipid peroxidation (LPO) in both testes and sperm compared to the Se-sufficient mice.|||
|Mouse||Se-deficient (0.02 ppm)
Se-excess (0.2 ppm); yeast-based Se.
Mice were fed for 4 months
|Se concentration and GPX activity (in testis) were significantly reduced. The fertility percentage and size of litter were both reduced in Se-deficient group.|||
|Aged mice||Inorganic Se 0.2 mg/kg body weight||Improved sperm parameters and increased expression of CatSper genes were observed in Se-treated group.|||
|Rabbit||Treated with Se nanoparticles (400 μg/kg) for 60 days||Improved serum testosterone levels were recorded in Se-treated group compared to the control. Besides, improved ejaculate volume and sperm quality parameters such as, sperm morphology, viability were observed.|||
|Ram||0.5 ppm organic Se; 0.2 ppm organic Se||A significantly higher concentration of Se and improved ejaculate and sperm quality were observed in seminal plasma of rams exposed to a feed containing 0.5 ppm organic Se compared to those who received 0.2 ppm organic Se.|||
(0.2 mg per kg);
(0.2 mg per kg)
|Ejaculate quality and sperm parameters were significantly improved in boars following dietary supplementation of organic Se (0.2 mg per kg) compared to those treated with sodium selenite at the same dose.|||
|Aardi buck||Sodium selenite 0.1 mg/kg,
Sodium selenite 0.05 mg/kg
|Improved sperm count and motility was observed in both Se-treated groups. However, relatively better outcomes were observed in 0.1 mg/kg group.|||
|Boar||0.5 ppm organic Se||Following 11 weeks of feeding trail, organic Se supplementation increased glutathione peroxidase 4 (GPX4) activity (raw semen) and number of seminal doses in boars.|||
|Boar||0.3 ppm organic Se;
0.3 ppm inorganic Se
|Following 12 weeks of Se supplementation, Se content and GPX activity were increased in semen of boars treated with organic and inorganic Se. Besides, semen quality parameters namely semen concentration and progressive motility of sperm were improved compared to the control group without Se. Improved resistance of liquid stored semen to hypo-osmotic shock and thermal tests, and improved fertility rates were observed in semen of boars treated with Se. All mentioned indices were slightly higher in the organic Se group compared to the inorganic group.|||
|Buffalo bulls||10 mg organic Se/animal twice a week;
10 mg inorganic Se/animal twice a week
|Three months long Se supplementation significantly improved the sperm quality parameters (ejaculate volume, sperm motility, concentration, and morphology) in buffalo bulls. Besides, testosterone concentrations were also increased in Se-treated groups.|||
|Saanen bucks||Inorganic Se 0.34 mg/kg body weight supplemented at ten-day intervals for three months||Se supplementation improved the testicular biometry and sperm parameters. GPX activity, plasma testosterone and LH levels significantly increased in Se-treated group from days 40 to 80 compared to the control group. These indices reached peak reached peak at day 80 of the trial.|||
|Bovine bull||In vitro fertilization (IVF) medium supplemented with Se (100 ng/mL)||A significant increase in sperm mitochondrial activity was observed after 1 h of incubation in Se-supplemented IVF medium. Moreover, Se supplementation after 2 h of incubation showed an increase in HOST-positive (hypo-osmotic swelling test) sperm and sperm acrosome integrity. Increased number of sperm bound to zona pellucida (ZP) was observed in Se-treated group compared to the control.|||
Selenium in Seminal Plasma and its Implication in Male Fertility
3.3. Combinatorial Effects of Se (as a Part of Micronutrient Supplement) on Male Fertility Outcomes (Animal Studies)
|Animal Model and Number||Treatment Regime and Duration||Key Findings||Ref.|
|Male CD-1 mice
(n = 12 per experimental group)
|Fertilix® (CellOxess, Princeton, NJ, USA) was supplemented for two months.
(Se 55 μg, zinc 7.5–11 mg,
Full spectrum natural vitamin E 104–290 mg,
Lycopene 7.5–15 mg
Carnitine blend 200–800 mg
Folic acid 400–500 mg
Vitamin C 30–90 mg).
|Eight weeks long pretreatment with the antioxidant formulation completely protected oxidative stress-induced DNA damage in Gpx5 KO mice sperm. In mouse models of scrotal heat stress, only 35% (19/54) of female mice became pregnant resulting in 169 fetuses with 18% fetal resorption (30/169). Conversely, in antioxidant pretreated group 74% (42/57) of female mice became pregnant, resulting in 427 fetuses with 9% fetal resorption (38/427).|||
|Four infertile male dogs with low blood Se levels (86.0–165.0 μg/L)||Organic Se 0.6 mg/kg and vitamin E (5 mg/kg) orally supplemented for 60 days.||Treated dogs showed improved sperm parameters. Increase in blood Se concentration (401 μg/L) was observed at the end of trial. When these dogs were used for matting purpose, bitches successfully conceived and gave birth to 4–6 pups.|||
|Sixteen healthy normospermic dogs (two patients were excluded after adaptation period)||A supplement comprising of Se 0.27 mg/kg vitamin E 250 mg/kg, vitamin B9 1.5 mg/kg, zinc 180 mg/kg, and n-3 PUFA 0.5%, given for 90 days.||In treated group, sperm quality parameters i.e., total sperm count, concentration, sperm vitality and membrane integrity were significantly improved compared to the control group.|||
3.4. Selenium and Sperm Cryopreservation
The entry is from 10.3390/antiox8080268
- Oldfield, J.E. The two faces of selenium. J. Nutr. 1987, 117, 2002–2008.
- Schwarz, K.; Foltz, C.M. Selenium as an integral part of factor 3 against dietary necrotic liver degeneration. J. Am. Chem. Soc. 1957, 79, 3292–3293.
- O’Dell, B.L.; Sunde, R.A. Handbook of Nutritionally Essential Mineral Elements; CRC Press: New York, NY, USA; Basel, Switzerland; Hong Kong, China, 1997.
- Ceko, M.J.; O’leary, S.; Harris, H.H.; Hummitzsch, K.; Rodgers, R.J. Trace elements in ovaries: Measurement and physiology. Biol. Reprod. 2016, 94, 86.
- Xiong, X.; Lan, D.; Li, J.; Lin, Y.; Li, M. Selenium supplementation during in vitro maturation enhances meiosis and developmental capacity of yak oocytes. Anim. Sci. J. 2018, 89, 298–306.
- Hedaoo, M.; Khllare, K.; Meshram, M.; Sahatpure, S.; Patil, M. Study of some serum trace minerals in cyclic and non-cyclic surti buffaloes. Vet. World 2008, 1, 71.
- Mirone, M.; Giannetta, E.; Isidori, A. Selenium and reproductive function. A systematic review. J. Endocrinol. Investig. 2013, 36, 28–36.
- Pappas, A.; Zoidis, E.; Surai, P.; Zervas, G. Selenoproteins and maternal nutrition. Comp. Biochem. Physiol. Part B Biochem. Mol. Biol. 2008, 151, 361–372.
- Foresta, C.; Flohé, L.; Garolla, A.; Roveri, A.; Ursini, F.; Maiorino, M. Male fertility is linked to the selenoprotein phospholipid hydroperoxide glutathione peroxidase. Biol. Reprod. 2002, 67, 967–971.
- Kommisrud, E.; Østerås, O.; Vatn, T. Blood selenium associated with health and fertility in Norwegian dairy herds. Acta Vet. Scand. 2005, 46, 229.
- Behne, D.; Höfer, T.; von Berswordt-Wallrabe, R.; Elger, W. Selenium in the testis of the rat: Studies on its regulation and its importance for the organism. J. Nutr. 1982, 112, 1682–1687.
- Behne, D.; Weiler, H.; Kyriakopoulos, A. Effects of selenium deficiency on testicular morphology and function in rats. J. Reprod. Fertil. 1996, 106, 291–297.
- Flohe, L. Selenium in mammalian spermiogenesis. Biol. Chem. 2007, 388, 987–995.
- Ahsan, U.; Kamran, Z.; Raza, I.; Ahmad, S.; Babar, W.; Riaz, M.; Iqbal, Z. Role of selenium in male reproduction—A review. Anim. Reprod. Sci. 2014, 146, 55–62.
- Irvine, D.S. Glutathione as a treatment for male infertility. Rev. Reprod. 1996, 1, 6–12.
- Conrad, M.; Schweizer, U. Mouse Models that Target Individual Selenoproteins. In Selenium: Its Molecular Biology and Role in Human Health; Hatfield, D.L., Schweizer, U., Tsuji, P.A., Gladyshev, V.N., Eds.; Springer: New York, NY, USA, 2016; pp. 567–578.
- Riese, C.; Michaelis, M.; Mentrup, B.; Gotz, F.; Kohrle, J.; Schweizer, U.; Schomburg, L. Selenium-dependent pre-and posttranscriptional mechanisms are responsible for sexual dimorphic expression of selenoproteins in murine tissues. Endocrinology 2006, 147, 5883–5892.
- Beckett, G.J.; Arthur, J.R. Selenium and endocrine systems. J. Endocrinol. 2005, 184, 455–465.
- Knapen, M.F.; Zusterzeel, P.L.; Peters, W.H.; Steegers, E.A. Glutathione and glutathione-related enzymes in reproduction: A review. Eur. J. Obstet. Gynecol. Reprod. Biol. 1999, 82, 171–184.
- Ursini, F.; Heim, S.; Kiess, M.; Maiorino, M.; Roveri, A.; Wissing, J.; Flohé, L. Dual function of the selenoprotein PHGPx during sperm maturation. Science 1999, 285, 1393–1396.
- Safarinejad, M.R.; Safarinejad, S. Efficacy of selenium and/or N-acetyl-cysteine for improving semen parameters in infertile men: A double-blind, placebo controlled, randomized study. J. Urol. 2009, 181, 741–751.
- Imai, H.; Hirao, F.; Sakamoto, T.; Sekine, K.; Mizukura, Y.; Saito, M.; Kitamoto, T.; Hayasaka, M.; Hanaoka, K.; Nakagawa, Y. Early embryonic lethality caused by targeted disruption of the mouse PHGPx gene. Biochem. Biophys. Res. Commun. 2003, 305, 278–286.
- Conrad, M.; Moreno, S.; Sinowatz, F.; Ursini, F.; Kölle, S.; Roveri, A.; Brielmeier, M.; Wurst, W.; Maiorino, M.; Bornkamm, G. The nuclear form of phospholipid hydroperoxide glutathione peroxidase is a protein thiol peroxidase contributing to sperm chromatin stability. Mol. Cell. Biol. 2005, 25, 7637–7644.
- Chabory, E.; Damon, C.; Lenoir, A.; Henry-Berger, J.; Vernet, P.; Cadet, R.; Saez, F.; Drevet, J. Mammalian glutathione peroxidases control acquisition and maintenance of spermatozoa integrity 1. J. Anim. Sci. 2010, 88, 1321–1331.
- Schneider, M.; Förster, H.; Boersma, A.; Seiler, A.; Wehnes, H.; Sinowatz, F.; Neumüller, C.; Deutsch, M.J.; Walch, A.; de Angelis, M.H. Mitochondrial glutathione peroxidase 4 disruption causes male infertility. FASEB J. 2009, 23, 3233–3242.
- Imai, H.; Suzuki, K.; Ishizaka, K.; Ichinose, S.; Oshima, H.; Okayasu, I.; Emoto, K.; Umeda, M.; Nakagawa, Y. Failure of the expression of phospholipid hydroperoxide glutathione peroxidase in the spermatozoa of human infertile males. Biol. Reprod. 2001, 64, 674–683.
- Imai, H.; Hakkaku, N.; Iwamoto, R.; Suzuki, J.; Suzuki, T.; Tajima, Y.; Konishi, K.; Minami, S.; Ichinose, S.; Ishizaka, K. Depletion of selenoprotein GPx4 in spermatocytes causes male infertility in mice. J. Biol. Chem. 2009, 284, 32522–32532.
- Meseguer, M.; Maria, J.; Simón, C.; Pellicer, A.; Remohí, J.; Garrido, N. Effect of sperm glutathione peroxidases 1 and 4 on embryo asymmetry and blastocyst quality in oocyte donation cycles. Fertil. Steril. 2006, 86, 1376–1385.
- Parillo, F.; Sylla, L.; Palombi, C.; Monaci, M.; Stradaioli, G. Immunocytochemical Localisation of Phospholipid Hydroperoxide Glutathione Peroxidase in Bull’s Spermatogenic Cells. Ital. J. Anim. Sci. 2014, 13, 3483.
- Olson, G.E.; Winfrey, V.P.; NagDas, S.K.; Hill, K.E.; Burk, R.F. Apolipoprotein E receptor-2 (ApoER2) mediates selenium uptake from selenoprotein P by the mouse testis. J. Biol. Chem. 2007, 282, 12290–12297.
- Hill, K.E.; Zhou, J.; McMahan, W.J.; Motley, A.K.; Atkins, J.F.; Gesteland, R.F.; Burk, R.F. Deletion of selenoprotein P alters distribution of selenium in the mouse. J. Biol. Chem. 2003, 278, 13640–13646.
- Kehr, S.; Malinouski, M.; Finney, L.; Vogt, S.; Labunskyy, V.M.; Kasaikina, M.V.; Carlson, B.A.; Zhou, Y.; Hatfield, D.L.; Gladyshev, V.N. X-ray fluorescence microscopy reveals the role of selenium in spermatogenesis. J. Mol. Biol. 2009, 389, 808–818.
- Noblanc, A.; Kocer, A.; Chabory, E.; Vernet, P.; Saez, F.; Cadet, R.; Conrad, M.; Drevet, J.R. Glutathione peroxidases at work on epididymal spermatozoa: An example of the dual effect of reactive oxygen species on mammalian male fertilizing ability. J. Androl. 2011, 32, 641–650.
- Rayman, M.P. Selenium and human health. Lancet 2012, 379, 1256–1268.
- Qazi, I.H.; Angel, C.; Yang, H.; Pan, B.; Zoidis, E.; Zeng, C.J.; Han, H.; Zhou, G.B. Selenium, Selenoproteins, and Female Reproduction: A Review. Molecules 2018, 23, 3053.
- Rayman, M.P. The importance of selenium to human health. Lancet 2000, 356, 233–241.
- Gladyshev, V.N.; Arnér, E.S.; Berry, M.J.; Brigelius-Flohé, R.; Bruford, E.A.; Burk, R.F.; Carlson, B.A.; Castellano, S.; Chavatte, L.; Conrad, M. Selenoprotein gene nomenclature. J. Biol. Chem. 2016, 291, 24036–24040.
- Zoidis, E.; Pappas, A. The health effects of selenoproteins. In Selenium: Sources, Functions and Health Effects; Aomori, C., Hokkaido, M., Eds.; Nova Science Publishers: New York, NY, USA, 2012.
- Boitani, C.; Puglisi, R. Selenium, a key element in spermatogenesis and male fertility. In Molecular Mechanisms in Spermatogenesis; Springer: Berlin/Heidelberg, Germany, 2009; pp. 65–73.
- Su, D.; Novoselov, S.V.; Sun, Q.-A.; Moustafa, M.E.; Zhou, Y.; Oko, R.; Hatfield, D.L.; Gladyshev, V.N. Mammalian selenoprotein thioredoxin-glutathione reductase roles in disulfide bond formation and sperm maturation. J. Biol. Chem. 2005, 280, 26491–26498.
- Michaelis, M.; Gralla, O.; Behrends, T.; Scharpf, M.; Endermann, T.; Rijntjes, E.; Pietschmann, N.; Hollenbach, B.; Schomburg, L. Selenoprotein P in seminal fluid is a novel biomarker of sperm quality. Biochem. Biophys. Res. Commun. 2014, 443, 905–910.
- Renko, K.; Werner, M.; Renner-Muller, I.; Cooper, T.G.; Yeung, C.H.; Hollenbach, B.; Scharpf, M.; Kohrle, J.; Schomburg, L.; Schweizer, U. Hepatic selenoprotein P (SePP) expression restores selenium transport and prevents infertility and motor-incoordination in Sepp-knockout mice. Biochem. J. 2008, 409, 741–749.
- Olson, G.E.; Winfrey, V.P.; NagDas, S.K.; Hill, K.E.; Burk, R.F. Selenoprotein P is required for mouse sperm development. Biol. Reprod. 2005, 73, 201–211.
- Turanov, A.A.; Malinouski, M.; Gladyshev, V.N. Selenium and male reproduction. In Selenium: Its Molecular Biology and Role in Human Health; Hatfield, D.L., Berry, M.J., Gladyshev, V.N., Eds.; Springer: New York, NY, USA, 2012; pp. 409–417.
- Kryukov, G.V.; Castellano, S.; Novoselov, S.V.; Lobanov, A.V.; Zehtab, O.; Guigó, R.; Gladyshev, V.N. Characterization of mammalian selenoproteomes. Science 2003, 300, 1439–1443.
- Behne, D.; Duk, M.; Elger, W. Selenium content and glutathione peroxidase activity in the testis of the maturing rat. J. Nutr. 1986, 116, 1442–1447.
- Nishimura, K.; Matsumiya, K.; Tsujimura, A.; Koga, M.; Kitamura, M.; Okuyama, A. Association of selenoprotein P with testosterone production in cultured Leydig cells. Arch. Androl. 2001, 47, 67–76.
- Koga, M.; Tanaka, H.; Yomogida, K.; Tsuchida, J.; Uchida, K.; Kitamura, M.; Sakoda, S.; Matsumiya, K.; Okuyama, A.; Nishimune, Y. Expression of selenoprotein-P messenger ribonucleic acid in the rat testis. Biol. Reprod. 1998, 58, 261–265.
- Oluboyo, A.; Adijeh, R.; Onyenekwe, C.; Oluboyo, B.; Mbaeri, T.; Odiegwu, C.; Chukwuma, G.; Onwuasoanya, U. Relationship between serum levels of testosterone, zinc and selenium in infertile males attending fertility clinic in Nnewi, south east Nigeria. Afr. J. Med. Med. Sci. 2012, 41, 51–54.
- Villaverde, A.I.S.; Fioratti, E.G.; Ramos, R.S.; Neves, R.C.; Ferreira, J.C.P.; Cardoso, G.S.; Padilha, P.M.; Lopes, M.D. Blood and seminal plasma concentrations of selenium, zinc and testosterone and their relationship to sperm quality and testicular biometry in domestic cats. Anim. Reprod. Sci. 2014, 150, 50–55.
- Richards, J.; Hallford, D.; Duff, G. Serum luteinizing hormone, testosterone, and thyroxine and growth responses of ram lambs fed locoweed (Oxvtropis sericea) and treated with vitamin e/selenium. Theriogenology 1999, 52, 1055–1066.
- Shi, L.; Zhang, C.; Yue, W.; Shi, L.; Zhu, X.; Lei, F. Short-term effect of dietary selenium-enriched yeast on semen parameters, antioxidant status and Se concentration in goat seminal plasma. Anim. Feed Sci. Technol. 2010, 157, 104–108.
- Ren, X.-M.; Wang, G.-G.; Xu, D.-Q.; Luo, K.; Liu, Y.-X.; Zhong, Y.-H.; Cai, Y.-Q. The protection of selenium on cadmium-induced inhibition of spermatogenesis via activating testosterone synthesis in mice. Food Chem. Toxicol. 2012, 50, 3521–3529.
- Shi, L.; Song, R.; Yao, X.; Ren, Y. Effects of selenium on the proliferation, apoptosis and testosterone production of sheep Leydig cells in vitro. Theriogenology 2017, 93, 24–32.
- Liu, H.; Li, X.; Qin, F.; Huang, K. Selenium suppresses oxidative-stress-enhanced vascular smooth muscle cell calcification by inhibiting the activation of the PI3K/AKT and ERK signaling pathways and endoplasmic reticulum stress. JBIC J. Biol. Inorg. Chem. 2014, 19, 375–388.
- Shi, L.; Song, R.; Yao, X.; Duan, Y.; Ren, Y.; Zhang, C.; Yue, W.; Lei, F. Effects of maternal dietary selenium (Se-enriched yeast) on testis development, testosterone level and testicular steroidogenesis-related gene expression of their male kids in Taihang Black Goats. Theriogenology 2018, 114, 95–102.
- Cerny, K.; Garbacik, S.; Skees, C.; Burris, W.; Matthews, J.; Bridges, P. Gestational form of selenium in free-choice mineral mixes affects transcriptome profiles of the neonatal calf testis, including those of steroidogenic and spermatogenic pathways. Biol. Trace Elem. Res. 2016, 169, 56–68.
- Brennan, K.M.; Burris, W.R.; Boling, J.A.; Matthews, J.C. Selenium content in blood fractions and liver of beef heifers is greater with a mix of inorganic/organic or organic versus inorganic supplemental selenium but the time required for maximal assimilation is tissue-specific. Biol. Trace Elem. Res. 2011, 144, 504–516.
- Liao, S.F.; Brown, K.R.; Stromberg, A.J.; Burris, W.R.; Boling, J.A.; Matthews, J.C. Dietary supplementation of selenium in inorganic and organic forms differentially and commonly alters blood and liver selenium concentrations and liver gene expression profiles of growing beef heifers. Biol. Trace Elem. Res. 2011, 140, 151–169.
- Watanabe, T.; Endo, A. Effects of selenium deficiency on sperm morphology and spermatocyte chromosomes in mice. Mutat. Res. Lett. 1991, 262, 93–99.
- Kaur, P.; Bansal, M.P. Effect of selenium-induced oxidative stress on the cell kinetics in testis and reproductive ability of male mice. Nutrition 2005, 21, 351–357.
- Adegoke, E.; Wang, X.; Wang, H.; Wang, C.; Zhang, H.; Zhang, G. Selenium (Na 2 SeO 3) Upregulates Expression of Immune Genes and Blood–Testis Barrier Constituent Proteins of Bovine Sertoli Cell In Vitro. Biol. Trace Elem. Res. 2018, 185, 332–343.
- Adegoke, E.; Xue, W.; Machebe, N.; Adeniran, S.; Hao, W.; Chen, W.; Han, Z.; Guixue, Z.; Peng, Z. Sodium Selenite inhibits mitophagy, downregulation and mislocalization of blood–testis barrier proteins of bovine Sertoli cell exposed to microcystin-leucine arginine (MC-LR) via TLR4/NF-kB and mitochondrial signaling pathways blockage. Ecotoxicol. Environ. Saf. 2018, 166, 165–175.
- Kaushal, N.; Bansal, M. Inhibition of CDC2/Cyclin B1 in response to selenium-induced oxidative stress during spermatogenesis: Potential role of Cdc25c and p21. Mol. Cell. Biochem. 2007, 298, 139–150.
- Kaushal, N.; Bansal, M. Selenium variation induced oxidative stress regulates p53 dependent germ cell apoptosis: Plausible involvement of HSP70-2. Eur. J. Nutr. 2009, 48, 221–227.
- Shalini, S.; Bansal, M. Alterations in selenium status influences reproductive potential of male mice by modulation of transcription factor NFκB. Biometals 2007, 20, 49–59.
- Shalini, S.; Bansal, M.P. Role of selenium in spermatogenesis: Differential expression of cjun and cfos in tubular cells of mice testis. Mol. Cell. Biochem. 2006, 292, 27–38.
- Shalini, S.; Bansal, M. Role of selenium in regulation of spermatogenesis: Involvement of activator protein 1. Biofactors 2005, 23, 151–162.
- Kaushal, N.; Bansal, M. Diminished reproductive potential of male mice in response to selenium-induced oxidative stress: Involvement of HSP70, HSP70-2, and MSJ-1. J. Biochem. Mol. Toxicol. 2009, 23, 125–136.
- Sattar, H.; Yang, J.; Zhao, X.; Cai, J.; Liu, Q.; Ishfaq, M.; Yang, Z.; Chen, M.; Zhang, Z.; Xu, S. Selenoprotein-U (SelU) knockdown triggers autophagy through PI3K–Akt–mTOR pathway inhibition in rooster Sertoli cells. Metallomics 2018, 10, 929–940.
- Bano, I.; Malhi, M.; Soomro, S.; Kandhro, S.; Awais, M.; Baloch, S.; Perveen, S.; Sajjad, H. Effect of Dietary Selenium Supplementation on Morphology and Antioxidant Status in Testes of Goat. J. Basic Appl. Sci. 2018, 14, 53–61.
- Stefanov, R.; Chervenkov, M.; Anev, G.; Maksimović, N.; Andreeva, M.; Ivanova, T.; Milovanović, A. Effect of supplementation with inorganic and organic selenium on sperm quality and quantity in north-east Bulgarian merino rams. Biotechnol. Anim. Husb. 2018, 34, 69–81.
- Asri-Rezaei, S.; Nourian, A.; Shalizar-Jalali, A.; Najafi, G.; Nazarizadeh, A.; Koohestani, M.; Karimi, A. Selenium supplementation in the form of selenium nanoparticles and selenite sodium improves mature male mice reproductive performances. Iran. J. Basic Med Sci. 2018, 21, 577.
- Martins, S.M.M.K.; Afonso, E.R.; Parazzi, L.J.; Andrade, A.F.C.D.; Leal, D.F.; Gameiro, A.H.; Moretti, A.D.S.A.; Arruda, R.P.D. Organic selenium supplementation is cost-effective for increasing the number of seminal doses produced by sexually mature boars. Rev. Bras. Zootec. 2018, 47.
- Liu, L.; He, Y.; Xiao, Z.; Tao, W.; Zhu, J.; Wang, B.; Liu, Z.; Wang, M. Effects of selenium nanoparticles on reproductive performance of male Sprague-Dawley rats at supranutritional and nonlethal levels. Biol. Trace Elem. Res. 2017, 180, 81–89.
- Zhou, J.-C.; Zheng, S.; Mo, J.; Liang, X.; Xu, Y.; Zhang, H.; Gong, C.; Liu, X.-L.; Lei, X.G. Dietary selenium deficiency or excess reduces sperm quality and testicular mRNA abundance of nuclear glutathione peroxidase 4 in rats. J. Nutr. 2017, 147, 1947–1953.
- Ibrahim, H.A.; Zhu, Y.; Wu, C.; Lu, C.; Ezekwe, M.O.; Liao, S.F.; Haung, K. Selenium-enriched probiotics improves murine male fertility compromised by high fat diet. Biol. Trace Elem. Res. 2012, 147, 251–260.
- Shalini, S.; Bansal, M. Dietary selenium deficiency as well as excess supplementation induces multiple defects in mouse epididymal spermatozoa: Understanding the role of selenium in male fertility. Int. J. Androl. 2008, 31, 438–449.
- Sánchez-Gutiérrez, M.; García-Montalvo, E.; Izquierdo-Vega, J.; Del Razo, L. Effect of dietary selenium deficiency on the in vitro fertilizing ability of mice spermatozoa. Cell Biol. Toxicol. 2008, 24, 321–329.
- Mohammadi, S.; Movahedin, M.; Mowla, S.J. Up-regulation of CatSper genes family by selenium. Reprod. Biol. Endocrinol. 2009, 7, 126.
- Abdel-Wareth, A.; Ahmed, A.; Hassan, H.; El-Sadek, M.A.; Ghazalah, A.; Lohakare, J. Nutritional impact of nano-selenium, garlic oil, and their combination on growth and reproductive performance of male Californian rabbits. Anim. Feed Sci. Technol. 2019, 249, 37–45.
- Baiomy, A.; Mohamed, A.; Mottelib, A. Effect of dietary selenium and vitamin E supplementation on productive and reproductive performance in rams. In Proceedings of the 14th International Congress of the International Society for Animal Hygiene (ISAH), Vechta, Germany, 9–23 July 2009; pp. 43–46.
- Jacyno, E.; Kawecka, M.; Kamyczek, M. Influence of inorganic Se+ vitamin E and organic Se+ vitamin E on reproductive performance of young boars. Agric. Food Sci. 2002, 11, 175–184.
- Hajalshaikh, A.; Al-Hassan, M.; Mohamed, H. The influence of injectable sodium selenite on semen characteristics and testosterone concentration in Aardi goats. Indian J. Anim. Res. 2015, 49, 793–797.
- Martins, S.M.M.K.; De Andrade, A.; Zaffalon, F.G.; Parazzi, L.J.; Bressan, F.F.; Pugine, S.M.P.; Melo, M.P.d.; Chiaratti, M.; Marino, C.T.; Afonso, E.R. Organic selenium increases PHGPx, but does not affect quality sperm in raw boar semen. Livest. Sci. 2014, 164, 175–178.
- Petrujkić, B.; Šefer, D.; Jovanović, I.; Jovičin, M.; Janković, S.; Jakovljević, G.; Beier, R.; Anderson, R. Effects of commercial selenium products on glutathione peroxidase activity and semen quality in stud boars. Anim. Feed Sci. Technol. 2014, 197, 194–205.
- El-Sharawy, M.; Eid, E.; Darwish, S.; Abdel-Razek, I.; Islam, M.R.; Kubota, K.; Yamauchi, N.; El-Shamaa, I. Effect of organic and inorganic selenium supplementation on semen quality and blood enzymes in buffalo bulls. Anim. Sci. J. 2017, 88, 999–1005.
- Lukusa, K.; Lehloenya, K. Selenium supplementation improves testicular characteristics and semen quality of Saanen bucks. Small Rumin. Res. 2017, 151, 52–58.
- Anchordoquy, J.P.; Anchordoquy, J.M.; Lizarraga, R.M.; Nikoloff, N.; Pascua, A.M.; Furnus, C.C. The importance of trace minerals copper, manganese, selenium and zinc in bovine sperm–zona pellucida binding. Zygote 2019, 1–8.
- Roychoudhury, S.; Sharma, R.; Sikka, S.; Agarwal, A. Diagnostic application of total antioxidant capacity in seminal plasma to assess oxidative stress in male factor infertility. J. Assist. Reprod. Genet. 2016, 33, 627–635.
- Gharagozloo, P.; Gutiérrez-Adán, A.; Champroux, A.; Noblanc, A.; Kocer, A.; Calle, A.; Pérez-Cerezales, S.; Pericuesta, E.; Polhemus, A.; Moazamian, A. A novel antioxidant formulation designed to treat male infertility associated with oxidative stress: Promising preclinical evidence from animal models. Hum. Reprod. 2016, 31, 252–262.
- Anel-López, L.; Ortega-Ferrusola, C.; Martínez-Rodríguez, C.; Álvarez, M.; Borragán, S.; Chamorro, C.; Peña, F.; Anel, L.; de Paz, P. Analysis of seminal plasma from brown bear (Ursus arctos) during the breeding season: Its relationship with testosterone levels. PLoS ONE 2017, 12, e0181776.
- Bertelsmann, H.; Keppler, S.; Höltershinken, M.; Bollwein, H.; Behne, D.; Alber, D.; Bukalis, G.; Kyriakopoulos, A.; Sieme, H. Selenium in blood, semen, seminal plasma and spermatozoa of stallions and its relationship to sperm quality. Reprod. Fertil. Dev. 2010, 22, 886–891.
- Hawkes, W.C.; Turek, P.J. Effects of dietary selenium on sperm motility in healthy men. J. Androl. 2001, 22, 764–772.
- Domosławska, A.; Zdunczyk, S.; Franczyk, M.; Kankofer, M.; Janowski, T. Selenium and vitamin E supplementation enhances the antioxidant status of spermatozoa and improves semen quality in male dogs with lowered fertility. Andrologia 2018, e13023.
- Domosławska, A.; Zduńczyk, S.; Niżański, W.; Jurczak, A.; Janowski, T. Effect of selenium and vitamin E supplementation on semen quality in dogs with lowered fertility. Bull. Vet. Inst. Pulawy 2015, 59, 85–90.
- Butt, M.A.; Shahid, M.Q.; Bhatti, J.A.; Khalique, A. Effect of Dietary Vitamin E and Selenium Supplementation on Physiological Responses and Reproductive Performance in Holstein Friesian Bulls during Humid Hot Summer. Pak. Vet. J. 2019, 1–5.
- Ghorbani, A.; Moeini, M.M.; Souri, M.; Hajarian, H. Influences of dietary selenium, zinc and their combination on semen characteristics and testosterone concentration in mature rams during breeding season. J. Appl. Anim. Res. 2018, 46, 813–819.
- Kirchhoff, K.; Failing, K.; Goericke-Pesch, S. Effect of dietary vitamin E and selenium supplementation on semen quality in Cairn Terriers with normospermia. Reprod. Domest. Anim. 2017, 52, 945–952.
- Domosławska, A.; Zduńczyk, S.; Janowski, T. Improvement of sperm motility within one month under selenium and vitamin E supplementation in four infertile dogs with low selenium status. J. Vet. Res. 2019.
- Alonge, S.; Melandri, M.; Leoci, R.; Lacalandra, G.M.; Caira, M.; Aiudi, G.G. The Effect of Dietary Supplementation of Vitamin E, Selenium, Zinc, Folic Acid, and N-3 Polyunsaturated Fatty Acids on Sperm Motility and Membrane Properties in Dogs. Animals 2019, 9, 34.
- Ran, M.-X.; Li, Y.; Zhang, Y.; Liang, K.; Ren, Y.-N.; Zhang, M.; Zhou, G.-B.; Zhou, Y.-M.; Wu, K.; Wang, C.-D. Transcriptome Sequencing Reveals the Differentially Expressed lncRNAs and mRNAs Involved in Cryoinjuries in Frozen-Thawed Giant Panda (Ailuropoda melanoleuca) Sperm. Int. J. Mol. Sci. 2018, 19, 3066.
- Dai, D.-H.; Qazi, I.H.; Ran, M.-X.; Liang, K.; Zhang, Y.; Zhang, M.; Zhou, G.-B.; Angel, C.; Zeng, C.-J. Exploration of miRNA and mRNA Profiles in Fresh and Frozen-Thawed Boar Sperm by Transcriptome and Small RNA Sequencing. Int. J. Mol. Sci. 2019, 20, 802.
- Jamali, N.U.; Kaka, A.; Khatri, P.; Malhi, M.; Naeem, M.; Memon, A.A.; Kaleri, R.R.; Janyaro, H.; Kalhoro, D.H. Effect of in vitro Selenium Addition to the Semen Extender on the Spermatozoa Characteristics before and after Freezing in Kundhi Buffalo Bull and in vivo Fertility Rate. Pak. J. Zool. 2019, 51, 317–323.
- Khalil, W.A.; El-Harairy, M.A.; Zeidan, A.E.; Hassan, M.A. Impact of selenium nano-particles in semen extender on bull sperm quality after cryopreservation. Theriogenology 2019, 126, 121–127.