The Sericin in semen cryopreservation: A review of its potential and challenges https://doi.org/10.12982/VIS.2026.019
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Abstract
Semen cryopreservation plays a critical role in animal reproduction, but oxidative stress and reactive oxygen species (ROS) generation during freezing and thawing can significantly reduce sperm quality. Sericin, a silk-derived biopolymer, has gained attention as a promising cryoprotectant due to its potent antioxidant, radical-scavenging, and antibacterial properties. This review explores sericin’s potential to improve sperm motility, viability, acrosome integrity, and DNA integrity across 12 species, including goats, bulls, buffalo, poultry, rabbits, boars, dogs, mice, horse, sheep, elephant, and humans by synthesizing findings from over 60 studies that utilized six different cryopreservation methodologies. Sericin has been shown to protect sperm during cryopreservation by reducing oxidative stress, preserving membrane integrity, and enhancing fertilizing capacity. Studies suggest that sericin supplementation may provide comparable or even superior protection compared to traditional cryoprotectants. Additionally, sericin can work synergistically with other additives, such as trehalose and glutathione, to further improve sperm motility and overall post-thaw quality. Despite these promising findings, several challenges hinder the widespread commercial application of sericin such as variability in composition, and scalability remain key limitations. Moreover, excessive sericin concentrations can lead to cytotoxicity, protein aggregation, osmotic imbalance, and increased viscosity, all of which may induce oxidative stress and ultimately impair sperm quality. Therefore, optimizing sericin concentrations is essential to maximize its cryoprotective benefits while minimizing potential adverse effects. Future research should focus on elucidating the molecular mechanisms underlying sericin’s protective effects and conducting field trials to validate its efficacy in diverse reproductive contexts. With further development, sericin holds great potential for advancing semen preservation and enhancing reproductive technologies.
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References
Aad, R., Dragojlov, I., Vesentini, S., 2024. sericin protein: structure, properties, and applications. J. Funct. Biomater. 15(11), 1-36.
Aghaz, F., Hajarian, H., KaramiShabankareh, H., 2016. In vitro culture medium (IVC) supplementation with sericin improves developmental competence of ovine zygotes. Reprod. Biol. 16(1), 87–90.
Aghaz, F., Khazaei, M., Vaisi-Raygani, A., Bakhtiyari, M., 2020. Cryoprotective effect of sericin supplementation in freezing and thawing media on the outcome of cryopreservation in human sperm. Aging. Male. 23(5), 469–476.
Aitken, R.J., Buckingham, D., Harkiss, D., 1993. Use of a xanthine oxidase free radical generating system to investigate the cytotoxic effects of reactive oxygen species on human spermatozoa. J. Reprod. Fertil. 97, 441–450.
Alvarez, J.G., Storey, B.T., 1989. Role of glutathione peroxidase in protecting mammalian spermatozoa from loss of motility caused by spontaneous lipid peroxidation. Gamete Res. 23(1), 77–90.
Appiah, M.O., Li, W., Zhao, J., Liu, H., Dong, Y., Xiang, J., Lu, W., 2020. Quercetin supplemented casein-based extender improves the post-thaw quality of rooster semen. Cryobiology. 94, 57-65.
Aramwit, P., Damrongsakkul, S., Kanokpanont, S., Srichana, T., 2010. Properties and antityrosinase activity of sericin from various extraction methods. Biotechnol. Appl. Biochem. 55, 91–98.
Aramwit, P., Sangcakul, A., 2007. The effects of sericin cream on wound healing in rats. Biosci. Biotechnol. Biochem. 71(10), 2473–2477.
Aramwit, P., Siritientong, T., Srichana, T., 2012. Potential applications of silk sericin, a natural protein from textile industry by-products. Waste. Manag. Res. 30(3), 217–224.
Asadi, Z., Safari-Faramani, R., Aghaz, F., 2023. Effects of adding antioxidant nanoparticles on sperm parameters of non-human species after the freezing and thawing process: a systematic review and meta-analysis. Anim. Reprod. Sci. 257, 1-14.
Atessahin, A., Bucak, M.N., Tuncer, P.B., Kizil, M., 2008. Effects of antioxidant additives on microscopic and oxidative parameters of Angora goat semen following the freeze-thawing process. Small. Rumin. Res. 77(1), 38-44.
Awda, B.J., Mackenzie-Bell, M., Buhr, M.M., 2009. Reactive oxygen species and boar sperm function. Biol. Reprod. 81(3), 553–561.
Baumber, J., Ball, B.A., Linfor, J.J., 2005. Assessment of the cryopreservation of equine spermatozoa in the presence of enzyme scavengers and antioxidants. Am. J. Vet. Res. 66(5), 772–779.
Bergeron, A., Brindle, Y., Blondin, P., Manjunath, P., 2007. Milk caseins decrease the binding of the major bovine seminal plasma proteins to sperm and prevent lipid loss from the sperm membrane during sperm storage. Biol. Reprod. 77(1), 120-126.
Bergeron, A., Crête, M. H., Brindle, Y., Manjunath, P., 2004. Low-density lipoprotein fraction from hen’s egg yolk decreases the binding of the major proteins of bovine seminal plasma to sperm and prevents lipid efflux from the sperm membrane. Biol. Reprod. 70, 708–717.
Bucak, M.N., Ateşşahin, A., Varışlı, Ö., Yüce, A., Tekin, N., Akçay, A., 2007. The influence of trehalose, taurine, cysteamine, and hyaluronan on ram semen: microscopic and oxidative stress parameters after freeze–thawing process. Theriogenology. 67(5), 1060-1067.
Cao, T.T., Zhang, Y.Q., 2017. The potential of silk sericin protein as a serum substitute or an additive in cell culture and cryopreservation. Amino. Acids. 49, 1029–1039.
Chaturvedi, D., Dhami, A.J., Chaudhari, D.V., 2022. Fortification of tris extender with mifepristone, sericin and taurine improves velocity and kinematics of fresh and frozen-thawed bovine spermatozoa. Indian J. Anim. Res. 56(3), 255–262.
Chohan, K.R., Griffin, J.T., Carrell, D.T., 2004. Evaluation of chromatin integrity in human sperm using acridine orange staining with different fixatives and after cryopreservation. Andrologia. 36(5), 321–326.
Darin-Bennett, A., White, I.G., 1977. Influence of the cholesterol content of mammalian spermatozoa on susceptibility to cold-shock. Cryobiology. 14, 466–470.
Das, G., Shin, H.S., Campos, E.V.R., Fraceto, L.F., Del Pilar Rodriguez-Torres, M., Mariano, K.C.F., De Araujo, D.R., Fernández-Luqueño, F., Grillo, R., Patra, J.K., 2021. Sericin-based nanoformulations: A comprehensive review on molecular mechanisms of interaction with organisms to biological applications. J. Nanobiotechnol. 19(30), 1-22.
Davis, B.K., 1981. Timing of fertilization in mammals: sperm cholesterol/phospholipid ratio as a determinant of the capacitation interval. Cell Biol. 78, 7560-7564.
Demra, R., Gandotra, V.K., Singh, A.K., Kumar, A., 2017. Validation of Sericin concentration in semen freezing media and its effect on post-thaw quality of buffalo bull semen. Vet. Pract. 18(1), 42–44.
Do, L.T.K., Namula, Z., Luu, V.V., Sato, Y., Taniguchi, M., Isobe, T., Otoi, T., 2014. Effect of sericin supplementation during in vitro maturation on the maturation, fertilization and development of porcine oocytes. Reprod. Dom. Anim. 49(2), e17–e20.
Dorji, P., Vongprolub, T., Pattarajinda, V., Chankitisakul, V., Chuawongboon, P., 2015. Effect of sericin supplement in Tris-egg yolk citrate extender on quality of cryopreserved semen from Thai native bulls. Bhutan J. Nat. Resour. Dev. 2(1), 11–18.
Drobnis, E.Z., Crowe, L.M., Berger, T., Anchordoguy, T.J., Overstreet, J.W., Crowe, J.H., 1993. Cold shock damage is due to lipid phase transitions in cell membrane: a demonstration using sperm as a model. J. Exp. Zool. 265, 432–437.
Dutta, M., Kadirvel, G., Borah, P., Sinha, S., Ahmed, K., Hazarika, G., Sharma, R., Choudhury, H., Deori, S., Das Gupta, M., Biswas, R.K., Tamuly, S., Barua, P.M., Hussain, J., 2023. Effect of membrane stabilizers on semen quality and sperm membrane protein expression during cryopreservation of goat semen. Cryo Lett. 44, 299–306.
Fatahian, R., Fatahian, A., Fatahian, E., Fatahian, H., 2021. A critical review on application of silk sericin and its mechanical properties in various industries. J. Res. Appl. Mech. Eng. 9(2), 1-11.
Ghasemi, M., Farshad, A., Hajarian, H., Banafshi, O., Asadollahi, V., Fathi, F., 2018. The effects of sericin on cryopreserved sperm cells and subsequent embryo development in mice. Int. J. Reprod. BioMed. 16(6), 405-412.
Gholami, D., Sharafi, M., Esmaeili, V., Nadri, T., Alaei, L., Riazi, G., Shahverdi, A., 2023. Beneficial effects of trehalose and gentiobiose on human sperm cryopreservation. PLoS One. 18(4), 1-22.
Habiba, E.S., Harby, S.A., El-Sayed, N.S., Omar, E.M., Bakr, B.A., Augustyniak, M., Hassan, M.A., 2023. Sericin and melatonin mitigate diethylnitrosamine-instigated testicular impairment in mice: Implications of oxidative stress, spermatogenesis, steroidogenesis, and modulation of Nrf2/WT1/SF-1 signaling pathways. Life Sci. 334, 122220.
Hajarian, H., Aghaz, F., Shabankareh, H.K., 2017. Replacement of serum with sericin in in vitro maturation and culture media: Effects on embryonic developmental competence of Sanjabi sheep embryo during breeding season. Theriogenology. 92, 144–148.
Hassan, E., Magdy, S., Attaallah, A., Gaber, E., Mansour, O., Gomaa, R.A., El Wakil, A., 2022. Silk sericin alleviates aberrant photoperiod-induced alterations in testicular and adrenal steroidogenesis in adult mice. Reprod. Biol. Endocrinol. 20(1), 1-15.
Hezavehei, M., Sharafi, M., Kouchesfahani, H. M., Henkel, R., Agarwal, A., Esmaeili, V., Shahverdi, A., 2018. Sperm cryopreservation: a review on current molecular cryobiology and advanced approaches. Reprod. Biomed. Online. 37(3), 327-339.
Holt, W., 2000. Basic aspects of frozen storage of semen. Anim. Reprod. Sci. 62, 3–22
Hozbor, F., Ledesma, A., Manes, J., Rios, G.L., Kaiser, G., Cano, A., Alberio, R., 2016. Improve intra-uterine insemination in rabbits using ultra-high temperature skim milk as extender to keep semen at room temperature. Andrologia. 48(2), 231–234.
Ikeda, K., Oumi, Y., Ogawa, A., Sasaki, M., Yamada, H., Terada, S., 2010. Cryopreservative solution using sericin. In Cells and Culture: Proceedings of the 20th ESACT Meeting, Dresden, Germany, 17-20 June 2007, pp. 675-678.
Isobe, T., Ikebata, Y., Onitsuka, T., Do, L.T.K., Sato, Y., Taniguchi, M., Otoi, T., 2013. Cryopreservation for bovine embryos in serum-free freezing medium containing silk protein sericin. Cryobiology. 67(2), 184–187.
Isobe, T., Ikebata, Y., Onitsuka, T., Wittayarat, M., Sato, Y., Taniguchi, M., Otoi, T., 2012. Effect of sericin on preimplantation development of bovine embryos cultured individually. Theriogenology. 78(4), 747–752.
Jia, B., Memon, S., Liang, J., Lv, C., Hong, Q., Wu, G., Quan, G., 2021. Trehalose modifies the protein profile of ram spermatozoa during cryopreservation. Theriogenology. 171, 21–29.
Johinke, D., De Graaf, S., Bathgate, R., 2014. Quercetin reduces the in vitro production of H2O2 during chilled storage of rabbit spermatozoa. Anim. Reprod. Sci. 151, 208–219.
Johnson, L.A., Weitze, K.F., Fiser, P., Maxwell, W.M.C., 2000. Storage of boar semen. Anim. Reprod. Sci. 62(1–3), 143–172.
Kadirvel, G., Kumar, S., Kumaresan, A., 2009. Lipid peroxidation, mitochondrial membrane potential and DNA integrity of spermatozoa in relation to intracellular reactive oxygen species in liquid and frozen-thawed buffalo semen. Anim. Reprod. Sci. 114(1–3), 125–134.
Kamal, M.M., Alam, M.E., Das, S.K., Yeasmin, M.S., Ahmed, S., Rahman, M.A., Das, D.K., Gofur, M.R., 2023. Effects of glucose and trehalose on tris-citric acid-egg yolk-fructose diluents for semen cryopreservation in goat. J. Adv. Vet. Anim. Res. 10, 169–177.
Karimi‐Sabet, M.J., Khodaei‐Motlagh, M., Masoudi, R., Sharafi, M., 2024. Zinc oxide nanoparticles preserve the quality and fertility potential of rooster sperm during the cryopreservation process. Reprod. Domest. Anim. 59(4), 1-7.
Kato, N., Sato, S., Yamanaka, A., Yamada, H., Fuwa, N., Nomura, M., 1998. Silk protein, sericin, inhibits lipid peroxidation and tyrosinase activity. Biosci. Biotechnol. Biochem. 62(1), 145–147.
Khalil, W.A., El-Rais, M.S., Hegazy, M.M., Hassan, M.A., El-Raghi, A.A., El-Moghazy, M.M., 2024. The effect of metallic nanoparticles supplementation in semen extender on post-thaw quality and fertilizing ability of Egyptian buffalo (Bubalus bubalis) spermatozoa. Biol. Trace Elem. Res. 2024, 1–18.
Khalil, W.A., Ismail, A.A., Sindi, R.A., Abdel-Khalek, A.K.E., Yousif, A.I., Swelum, A.A., El-Harairy, M.A., 2023. Ethanolic purslane (Portulaca oleracea) leaf extract improves the quality of cryopreserved goat semen. Andrologia. 2023(1), 1-13.
Khatun, H., Egashira, J., Sakatani, M., Takenouchi, N., Tatemoto, H., Wada, Y., Yamanaka, K.I., 2018. Sericin enhances the developmental competence of heat‐stressed bovine embryos. Mol. Reprod. Dev. 85(8-9), 696–708.
Khatun, H., Yamanaka, K.I., Sugimura, S., 2024. Antioxidant sericin averts the disruption of oocyte–follicular cell communication triggered by oxidative stress. Mol. Hum. Reprod. 30(2), 1-12.
Khye, K.C., Yusuf, T.L., Satrio, F.A., Karja, N.W.K., 2021. Quality of chilled canine semen in tris-egg yolk extender supplemented with sericin. J. Kedokteran Hewan-Indonesian J. Vet. Sci. 15(1), 15–20.
Küçük, N., Raza, S., Matsumura, K., Uçan, U., Serin, İ., Ceylan, A., Aksoy, M., 2021. Effect of different carboxylated poly l-lysine and dimethyl sulfoxide combinations on post-thaw rabbit sperm functionality and fertility. Cryobiology. 102, 127–132.
Kumar, G., Yadav, D.K., Saxena, A., Sachan, V., Kumar, A., Swain, D.K., Yadav, B., 2023. Effect of trehalose on cryopreservation of Hariana bull spermatozoa. Indian J. Vet. Sci. Biotech. 19(6), 1–4.
Kumar, P., Kumar, D., Sikka, P., Singh, P., 2015. Sericin supplementation improves semen freezability of buffalo bulls by minimizing oxidative stress during cryopreservation. Anim. Reprod. Sci. 152, 26–31.
Kumar, S., Gupta, H.P., Prasad, S., Ambwani, T.K., Sharma, R.K., Singh, J.L., 2024. Studies on the quality and antioxidant status of cryopreserved Pantja buck semen supplemented with aqueous extract of rosemary and sericin in tris extender. Available online: https://doi.org/10.1089/bio.2024.008.
Kundu, S.C., Dash, B.C., Dash, R., Kaplan, D.L., 2008. Natural protective glue protein, sericin bioengineered by silkworms: Potential for biomedical and biotechnological applications. Prog. Polym. Sci. 33(10), 998–1012.
Kunz, R.I., Brancalhão, R.M.C., Ribeiro, L.D.F.C., Natali, M.R.M., 2016. Silkworm sericin: Properties and biomedical applications. BioMed Res. Int. 2016, 1–19.
Kweon, H., Yeo, J., Lee, K., Park, Y.H., Nahm, J.H., Cho, C., 2000. Effects of Poloxamer on the gelation of silk sericin. Macromol. Rapid Commun. 21(16), 1302–1305.
Lagares, M.A., Martins, H.S., Carvalho, I.A., Junior, C.A., Souza, M.R., Penna, C.F.A.M., Henry, M.R.J.M., 2012. Caseinate protects stallion sperm during semen cooling and freezing. CryoLetters. 33(3), 213-218.
Lamboni, L., Gauthier, M., Yang, G., Wang, Q., 2015. Silk sericin: a versatile material for tissue engineering and drug delivery. Biotechnol. Adv. 33(8), 1855-1867.
Lenzi, A., Lombardo, F., Gandini, L., Culasso, F., Dondero, F., 1992. Glutathione therapy for male infertility. Arch. Androl. 29(1), 65-68.
Li, Y.G., Ji, D.F., Lin, T.B., Zhong, S., Hu, G.Y., Chen, S., 2008. Protective effect of sericin peptide against alcohol-induced gastric injury in mice. Chin. Med. J. 121(20), 2083-2087.
Losano, J.D.A., Angrimani, D.S.R., Rui, B.R., Bicudo, L.C., Dalmazzo, A., Silva, B.C.S., Viana, C.H.C., Mendes, C.M., Assumpcao, M.E.O.A., Barnabee, V.H., Nichi, M., 2018. The addition of docosahexaenoic acid (DHA) and antioxidants (glutathione peroxidase and superoxide dismutase) in extenders to epididymal sperm cryopreservation in bulls. Zygote. 26(3), 199-206.
Lucio, C.D.F., Regazzi, F.M., Silva, L.C.G., Angrimani, D.D.S.R., Nichi, M., Vannucchi, C.I., 2016. Oxidative stress at different stages of two-step semen cryopreservation procedures in dogs. Theriogenology. 85(9), 1568-1575.
Mazur, P., 2004. Principles of cryobiology. In: Fuller, B.J., Lane, N., Benson, E.E. (Eds.), Life in the frozen state. CRC Press, Boca Raton, pp. 3–65.
Michaille, J.J., Garel, A., Prudhomme, J.C., 1989. The expression of five middle silk gland specific genes is territorially regulated during the larval development of Bombyx mori. Insect Biochem. 19(1), 19–27.
Miguel, G.A., Álvarez-López, C., 2020. Extraction and antioxidant activity of sericin, a protein from silk. Braz. J. Food Technol. 23, 1-14.
Miyamoto, Y., Oishi, K., Yukawa, H., Noguchi, H., Sasaki, M., Iwata, H., Hayashi, S., 2012. Cryopreservation of human adipose tissue-derived stem/progenitor cells using the silk protein sericin. Cell Transplant. 21(2-3), 617-622.
Mondal, M., Trivedy, K., Kumar, S.N., 2007. The silk proteins, sericin and fibroin in silkworm, Bombyx mori Linn. a review. Caspian J. Environ. Sci. 5(2), 63–76.
Muiño, R., Fernandez, M., Peña, A.I., 2007. Post‐thaw survival and longevity of bull spermatozoa frozen with an egg yolk‐based or two egg yolk‐free extenders after an equilibration period of 18 h. Reprod. Domest. Anim. 42(3), 305-311.
Nasirabadi, M.H., Shirazi, A., Kadivar, A., Shams-Esfandabadi, N., Mohebbi, A., Ahmadi, E., 2019. Sericin ameliorates the capacitation state and chromatin integrity of frozen-thawed stallion spermatozoa by reducing oxidative stress. Avicenna J. Med. Biotechnol. 11(3), 245-252.
Ohnishi, K., Murakami, M., Morikawa, M., Yamaguchi, A., 2012. Effect of the silk protein sericin on cryopreserved rat islets. J. Hepato Bil-Pan Sci. 19, 354–360.
Öztürk, A.E., Bodu, M., Bucak, M.N., Ağır, V., Özcan, A., Keskin, N., İli, P., Topraggaleh, T.R., Sidal, H., Başpınar, N., Dursun, Ş., 2020. The synergistic effect of trehalose and low concentrations of cryoprotectants can improve post-thaw ram sperm parameters. Cryobiology. 95, 157-163.
Park, K.S., Chun, J.L., Kim, E.Y., Lee, J.H., Lee, H.S., Yamauchi, N., LEE, K.B., Kim, M.K., 2018. Protective effect of trehalose on canine spermatozoa in cryopreservation. J. Fac. Agr. Kyushu Univ. 63(1), 53-60.
Parks, J.E., Graham, J.K., 1992. Effects of cryopreservation procedures on sperm membranes. Theriogenology. 38(2), 209–222.
Partyka, A., Niżański, W., 2021. Supplementation of avian semen extenders with antioxidants to improve semen quality—Is it an effective strategy? Antioxidants. 10(12), 1-17.
Patel, T.M., Dhami, A.J., Chaudhri, D.V., 2020. Correlations amongst functional and morphological attributes and oxidative markers of fresh and cryopreserved semen of Gir and Murrah bulls. Indian J. Vet. Sci. Biotechnol. 15(3), 24-29.
Patel, T.M., Dhami, A.J., Chaudhari, D.V., Pathan, M.M., 2019. Influence of antioxidant sericin in tris extender on oxidative markers during cryopreservation (–196ºC) of bovine semen. Indian J. Vet. Sci. Biotechnol. 15(2), 34-37.
Paul, D.R., Talukdar, D., Ahmed, F.A., Lalrintluanga, K., Kalita, G., Tolenkhomba, T.C., Chakravarty, H., Katiyar, R., Khargharia, G., Deori, S., 2024. Effect of selenium nanoparticles on the quality and fertility of short-term preserved boar semen. Front. Vet. Sci. 10, 1-7.
Rahal, A., Kumar, A., Singh, V., Yadav, B., Tiwari, R., Chakraborty, S., Dhama, K., 2014. Oxidative stress, prooxidants, and antioxidants: The interplay. Biomed Res Int. 2014, 1-19.
Ratchamak, R., Authaida, S., Boonkum, W., Chankitisakul, V., 2023. Improvement of rooster semen freezability and fertility rate after sericin supplementation in freezing semen extender. Anim. Biosci. 36(10), 1530-1535.
Ratchamak, R., Ratsiri, T., Kheawkanha, T., Vongpralub, T., Boonkum, W., Chankitisakul, V., 2020. Evaluation of cryopreserved boar semen after supplementation sericin from silkworm (Bombyx mori) in semen extender. Anim. Sci. J. 91(1), 1-7.
Raza, S., Uçan, U., Aksoy, M., Erdoğan, G., Ceylan, A., Serin, I., 2019. Silk protein sericin pretreatment enhances osmotic tolerance and post-thaw sperm quality but reduces the ability of sperm cells to undergo in vitro induced acrosome reaction in rabbit. Cryobiology. 90, 1-7.
Raza, S., Uçan, U., Aksoy, M., Erdoğan, G., Naseer, Z., Khan, K., 2024. Sericin-enriched rabbit semen preservation: implications for short-term storage quality and fertility at 4 or 15°C. Animals. 14(23), 1-11.
Reddy, V.S., Yadav, B., Yadav, C.L., Anand, M., Swain, D.K., Kumar, D., Yadav, S., 2018. Effect of sericin supplementation on heat shock protein 70 (HSP70) expression, redox status and post-thaw semen quality in goat. Cryobiology. 84, 33-39.
Saphungthong, B., Thammawung, S., Sirisathien, S., 2023. Cryoprotective effect of sericin on re-freezing of bull spermatozoa and subsequent embryonic development after in vitro fertilization. Vet. Integr. Sci. 21(1), 1-10.
Shaikh, K.Q., Suthar, H.N.B., Sutaria, P., Sharma, V., 2016. Comparison of fresh semen parameters with frozen-thawed semen following incorporation of trehalose. Life Sci. Leaflets. 76, 107-115.
Shaw, J.T.B., Smith, S.G., 1951. Amino-acids of silk sericin. Nature. 168(4278), 745.
Shin, S., Yeon, S., Park, D., Oh, J., Kang, H., Kim, S., Joo, S.S., Lim, W.T., Lee, J.Y., Choi, K.C., Kim, K.Y., Kim, S.U., Kim, J.C., Kim, Y.B., 2010. Silk amino acids improve physical stamina and male reproductive function of mice. Biol. Pharm. Bull. 33(2), 273-278.
Shu, W.H., Yang, S.H., Wei, M., Liu, X.C., Chen, Z.X., Wei, C.Y., Zhang, X., Si, L.N., Chen, Z.H., Qiao, Y.B., Li, D.M., Sun, T.C., Cheng, L.Y., 2023. Effects of sericin on oxidative stress and PI3K/AKT/mTOR signal pathway in cryopreserved mice ovarian tissue. Cryobiology. 111, 16-25.
Silva, A.S., Costa, E.C., Reis, S., Spencer, C., Calhelha, R.C., Miguel, S.P., Ribeiro, M.P., Barros, L., Vaz, J.A., Coutinho, P., 2022. Silk sericin: A promising sustainable biomaterial for biomedical and pharmaceutical applications. Polymers. 14(22), 1-22.
Silva, V.R., Ribani, M., Gimenes, M.L., Scheer, A.P., 2012. High molecular weight sericin obtained by high-temperature and ultrafiltration process. Procedia. Eng. 42, 833–841.
Sonseeda, P., Somtao, S., Laophaiboon, B., Duangjinda, M., Vongpralub, T., 2015. Effects of sericin supplementation on the sperm survival of cooled stored chicken semen. Khon. Kaen. Agr. J. 43(2), 103-106.
Takahashi, M., Tsujimoto, K., Yamada, H., Takagi, H., Nakamori, S., 2003. The silk protein, sericin, protects against cell death caused by acute serum deprivation in insect cell culture. Biotechnol. Lett. 25, 1805-1809.
Takechi, T., Wada, R., Fukuda, T., Harada, K., Takamura, H., 2014. Antioxidant activities of two sericin proteins extracted from cocoon of silkworm (Bombyx mori) measured by DPPH, chemiluminescence, ORAC and ESR methods. Biomed. Rep. 2(3), 364–369.
Terada, S., Nishimura, T., Sasaki, M., Yamada, H., Miki, M., 2002. Sericin, a protein derived from silkworms, accelerates the proliferation of several mammalian cell lines including a hybridoma. Cytotechnology. 40, 3-12.
Terada, S., Yanagihara, K., Kaito, K., Miki, M., Sasaki, M., Tsujimoto, K., 2005. Silk protein sericin accelerates proliferation of various mammalian cells. In Proceedings of the 18th ESACT Meeting Granada, Spain, 11–14 May 2003, pp. 585-587.
Theangeaw, K., Thongthip, N., Thongphakdee, A., Kiatsomboon, S., Noimoon, S., Angkawanish, T., Pinyopummin, A., 2023. Effect of sericin supplementation on post-thawed Asian elephant (Elephas maximus) semen quality. Kasetsart Univ. J. Sci. Technol. 12(3), 23-37.
Ugur, M.R., Saber Abdelrahman, A., Evans, H.C., Gilmore, A.A., Hitit, M., Arifiantini, R.I., Memili, E., 2019. Advances in cryopreservation of bull sperm. Front. Vet. Sci. 6, 268.
Ugur, M.R., Saber Abdelrahman, A., Evans, H.C., Gilmore, A.A., Hitit, M., Arifiantini, R.I., Memili, E., 2019. Advances in cryopreservation of bull sperm. Front. Vet. Sci. 6, 1-15.
Verdanova, M., Pytlik, R., Kalbacova, M.H., 2014. Evaluation of sericin as a fetal bovine serum-replacing cryoprotectant during freezing of human mesenchymal stromal cells and human osteoblast-like cells. Biopreserv. Biobank. 12(2), 99-105.
Vijyeta, H.P., Kavani, F.S., Vagh, A.A., Solanki, G.B., Vala, K.B., Singh, V.K., Dham, A.J., 2024. Effect of supplementation of antioxidant sericin in semen extender on post-thaw sperm quality, oxidative status, and fertility of Jaffarabadi buffalo bull semen. Indian J. Vet. Sci. Biotechnol. 20(2), 79-84.
Watson, P.F., 2000. The causes of reduced fertility with cryopreserved semen. Anim. Reprod. Sci. 60(3–4), 481–492.
White, I.G., 1993. Lipids and calcium uptake of sperm in relation to cold shock and preservation: a review. Reprod. Fertil. Dev. 5, 639-658.
Wu, J.H., Wang, Z., Xu, S.Y., 2007. Preparation and characterization of sericin powder extracted from silk industry wastewater. Food. Chem. 103(4), 1255-1262.
Xu, B., Wang, Z., Wang, R., Song, G., Zhang, Y., Su, R., Liu, Y., Li, J., Zhang, J., 2022. Metabolomics analysis of buck semen cryopreserved with trehalose. Front. Genet. 13, 1-11.
Yadava, C.L., Saxena, A., Yadav, B., Singh, V., Reddy, A.V.S., Patel, A., Kumar, A., Yadav, S., 2018. Effect of sericin supplementation on the semen quality of cryopreserved Hariana bull semen. Rumin. Sci. 7(1), 93-96.
Yanagihara, K., Terada, S., Miki, M., Sasaki, M., Yamada, H., 2006. Effect of the silk protein sericin on the production of adenovirus-based gene-therapy vectors. Biotechnol. Appl. Biochem. 45, 59–64.
Yangngam, Y., Chapanya, S., Vongpralub, T., Boonkum, W., Chankitisakul, V., 2021. Effect of semen extender supplementation with sericin on post-thaw dairy bull sperm quality and lipid peroxidation. Czech J. Anim. Sci. 66(1), 13-20.
Yasmin, C., Otoi, T., Setiadi, M., Karja, N., 2015. Maturation and fertilisation of sheep oocytes cultured in serum-free medium containing silk protein sericin. Acta Vet. Hung. 63(1), 110-117.
Younus, A.M., Yamanaka, T., Shimada, M., 2024. The protective effects of antioxidants against endogenous and exogenous oxidative stress on bull sperm. In Vitro Cell. Dev. Biol. Anim. 60; 969-982.
Zhang, W., Yu, Y., Qi, Y., Zhao, W., Zhang, H., Tao, C., Li, J., 2024. Effect of sericin on cryopreservation of semen of Chengde hornless black goat. Acta Vet. Zootech. Sinica. 55(10), 4466–4474.
Zhang, Y.Q., 2002. Applications of natural silk protein sericin in biomaterials. Biotechnol. Adv. 20(2), 91-100.
Zhao, J.G., Zhang, Y.Q., 2016. A new estimation of the total flavonoids in silkworm cocoon sericin layer through aglycone determination by hydrolysis-assisted extraction and HPLC-DAD analysis. Food Nutr. Res. 60(1), 1-8.
