{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T23:29:11Z","timestamp":1785281351309,"version":"3.55.0"},"reference-count":70,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2024,9,18]],"date-time":"2024-09-18T00:00:00Z","timestamp":1726617600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institute of Health","doi-asserted-by":"publisher","award":["5R33AT010637-01A1"],"award-info":[{"award-number":["5R33AT010637-01A1"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institute of Health","doi-asserted-by":"publisher","award":["3R33AT010637-02S1"],"award-info":[{"award-number":["3R33AT010637-02S1"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"The National Center for Complementary and Integrative Health","doi-asserted-by":"publisher","award":["5R33AT010637-01A1"],"award-info":[{"award-number":["5R33AT010637-01A1"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"The National Center for Complementary and Integrative Health","doi-asserted-by":"publisher","award":["3R33AT010637-02S1"],"award-info":[{"award-number":["3R33AT010637-02S1"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Malcolm Feist Endowed Chair in Diabetes","award":["5R33AT010637-01A1"],"award-info":[{"award-number":["5R33AT010637-01A1"]}]},{"name":"Malcolm Feist Endowed Chair in Diabetes","award":["3R33AT010637-02S1"],"award-info":[{"award-number":["3R33AT010637-02S1"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biomolecules"],"abstract":"<jats:p>Leydig cells are the primary source of testosterone or androgen production in male mammals. The blood\u2013testis barrier (BTB) maintains structural integrity and safeguards germ cells from harmful substances by blocking their entry into the seminiferous tubules. L-cysteine is essential to the production of glutathione, a powerful antioxidant crucial to protecting against oxidative stress-induced damage. Animal studies have demonstrated the protective effect of L-cysteine in preventing testicular damage caused by chemicals or radiation. This study examines whether L-cysteine enhances the expression of testosterone biosynthesis and the BTB genes in human Leydig cells and THP-1 monocytes. The Leydig cells and THP-1 monocytes were treated with L-cysteine for 24 h. RNA was extracted following treatment, and the gene expression was analyzed using quantitative RT-PCR. Testosterone levels in the cell supernatant were measured using an ELISA kit. L-cysteine treatment in Leydig cells significantly upregulated the expression of CYP11A1 (p = 0.03) and the BTB genes CLDN1 (p = 0.03), CLDN11 (p = 0.02), and TJP1 (p = 0.02). Similarly, L-cysteine significantly upregulated the expression of CYP11A1 (p = 0.03) and CYP19A1 (p &lt; 0.01), and the BTB genes CLDN1 (p = 0.04), CLDN2 (p &lt; 0.01), CLDN4 (p &lt; 0.01), CLDN11 (p &lt; 0.01), and TJP1 (p = 0.03) in THP-1 monocytes. Further, L-cysteine supplementation increased the testosterone secretion levels in human Leydig cells. The findings suggest that L-cysteine supplementation could be used as an adjuvant therapy to promote the integrity of the BTB genes, testosterone biosynthesis and secretion, and the maintenance of testicular functions, which in turn mitigates the risk of male infertility.<\/jats:p>","DOI":"10.3390\/biom14091171","type":"journal-article","created":{"date-parts":[[2024,9,18]],"date-time":"2024-09-18T09:43:13Z","timestamp":1726652593000},"page":"1171","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["L-Cysteine Upregulates Testosterone Biosynthesis and Blood\u2013Testis Barrier Genes in Cultured Human Leydig Cells and THP-1 Monocytes and Increases Testosterone Secretion in Human Leydig Cells"],"prefix":"10.3390","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7905-1887","authenticated-orcid":false,"given":"Jeffrey","family":"Justin Margret","sequence":"first","affiliation":[{"name":"Department of Pediatrics, Louisiana State University Health Sciences Center, Shreveport, LA 71103, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sushil K.","family":"Jain","sequence":"additional","affiliation":[{"name":"Department of Pediatrics, Louisiana State University Health Sciences Center, Shreveport, LA 71103, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2024,9,18]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"311","DOI":"10.5858\/2007-131-311-AILALC","article-title":"An in-depth look at Leydig cell tumor of the testis","volume":"131","author":"Axiotis","year":"2007","journal-title":"Arch. Pathol. Lab. Med."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"955","DOI":"10.1007\/s43032-021-00554-x","article-title":"Leydig Cell-Specific DAX1-Deleted Mice Has Higher Testosterone Level in the Testis during Pubertal Development","volume":"29","author":"Kumar","year":"2022","journal-title":"Reprod. Sci."},{"key":"ref_3","unstructured":"Aladamat, N., and Tadi, P. (2024). Histology, Leydig Cells, StatPearls Publishing."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1159\/000200077","article-title":"Fetal Leydig cell origin and development","volume":"3","author":"Griswold","year":"2009","journal-title":"Sex. Dev."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Marinkovic, D.Z., Medar, M.L.J., Becin, A.P., Andric, S.A., and Kostic, T.S. (2021). Growing up under Constant Light: A Challenge to the Endocrine Function of the Leydig Cells. Front. Endocrinol., 12.","DOI":"10.3389\/fendo.2021.653602"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1377","DOI":"10.1007\/s13258-023-01444-z","article-title":"VDR promotes testosterone synthesis in mouse Leydig cells via regulation of cholesterol side chain cleavage cytochrome P450 (Cyp11a1) expression","volume":"45","author":"Hu","year":"2023","journal-title":"Genes. Genom."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"3808","DOI":"10.1210\/jcem.87.8.8763","article-title":"Compound heterozygous mutations in the cholesterol side-chain cleavage enzyme gene (CYP11A) cause congenital adrenal insufficiency in humans","volume":"87","author":"Katsumata","year":"2002","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1177\/000992280104000708","article-title":"Lipoid congenital adrenal hyperplasia (CAH): Patient report and a mini-review","volume":"40","author":"Richmond","year":"2001","journal-title":"Clin. Pediatr."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.mce.2009.01.023","article-title":"Leydig cells: From stem cells to aging","volume":"306","author":"Chen","year":"2009","journal-title":"Mol. Cell Endocrinol."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Acikel-Elmas, M., Algilani, S.A., Sahin, B., Bingol Ozakpinar, O., Gecim, M., Koroglu, K., and Arbak, S. (2023). Apocynin Ameliorates Monosodium Glutamate Induced Testis Damage by Impaired Blood-Testis Barrier and Oxidative Stress Parameters. Life, 13.","DOI":"10.3390\/life13030822"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Luaces, J.P., Toro-Urrego, N., Otero-Losada, M., and Capani, F. (2023). What do we know about blood-testis barrier? current understanding of its structure and physiology. Front. Cell Dev. Biol., 11.","DOI":"10.3389\/fcell.2023.1114769"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Bhat, A.A., Syed, N., Therachiyil, L., Nisar, S., Hashem, S., Macha, M.A., Yadav, S.K., Krishnankutty, R., Muralitharan, S., and Al-Naemi, H. (2020). Claudin-1, A Double-Edged Sword in Cancer. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21020569"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"e13636","DOI":"10.1111\/and.13636","article-title":"Evaluation of blood-testis barrier integrity in terms of adhesion molecules in nonobstructive azoospermia","volume":"52","author":"Aydin","year":"2020","journal-title":"Andrologia"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"572","DOI":"10.4103\/1008-682X.125401","article-title":"Blood-testis barrier and spermatogenesis: Lessons from genetically-modified mice","volume":"16","author":"Jiang","year":"2014","journal-title":"Asian J. Androl."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"2612","DOI":"10.1210\/en.2007-1245","article-title":"Effect of glutathione depletion on Leydig cell steroidogenesis in young and old brown Norway rats","volume":"149","author":"Chen","year":"2008","journal-title":"Endocrinology"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1007\/s12011-018-1571-z","article-title":"Beneficial Effects of N-Acetyl-L-cysteine or Taurine Pre- or Post-treatments in the Heart, Spleen, Lung, and Testis of Hexavalent Chromium-Exposed Mice","volume":"190","author":"Bosgelmez","year":"2019","journal-title":"Biol. Trace Elem. Res."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.mrgentox.2015.09.004","article-title":"Ameliorative effects of N-acetylcysteine on fluoride-induced oxidative stress and DNA damage in male rats\u2019 testis","volume":"792","author":"Feng","year":"2015","journal-title":"Mutat. Res. Genet. Toxicol. Environ. Mutagen."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"319","DOI":"10.29271\/jcpsp.2023.03.319","article-title":"Protective Effects of N-Acetyl Cysteine on Undescended Testis after Orchiopexy: A Rat-model Study","volume":"33","author":"Kemahli","year":"2023","journal-title":"J. Coll. Physicians Surg. Pak."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1515\/hmbci-2022-0050","article-title":"The effect of co-administration of artemisinin and N-acetyl cysteine on antioxidant status, spermatological parameters and histopathology of testis in adult male mice","volume":"44","author":"Abedi","year":"2023","journal-title":"Horm. Mol. Biol. Clin. Investig."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1016\/j.jpedsurg.2019.09.028","article-title":"The long term effects of intrascrotal low dose and high dose N-acetylcysteine on testis damage in rat model of testicular torsion","volume":"55","author":"Mammadov","year":"2020","journal-title":"J. Pediatr. Surg."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1069","DOI":"10.1016\/j.ajem.2016.03.021","article-title":"Investigation of N-acetylcysteine on contralateral testis tissue injury by experimental testicular torsion: Long-term effect","volume":"34","author":"Bodur","year":"2016","journal-title":"Am. J. Emerg. Med."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Jannatifar, R., Parivar, K., Roodbari, N.H., and Nasr-Esfahani, M.H. (2019). Effects of N-acetyl-cysteine supplementation on sperm quality, chromatin integrity and level of oxidative stress in infertile men. Reprod. Biol. Endocrinol., 17.","DOI":"10.1186\/s12958-019-0468-9"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Monageng, E., Offor, U., Takalani, N.B., Mohlala, K., and Opuwari, C.S. (2023). A Review on the Impact of Oxidative Stress and Medicinal Plants on Leydig Cells. Antioxidants, 12.","DOI":"10.3390\/antiox12081559"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"2334","DOI":"10.1002\/jcb.24578","article-title":"L-cysteine and hydrogen sulfide increase PIP3 and AMPK\/PPARgamma expression and decrease ROS and vascular inflammation markers in high glucose treated human U937 monocytes","volume":"114","author":"Manna","year":"2013","journal-title":"J. Cell Biochem."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1007\/s11010-014-2009-3","article-title":"L-Cysteine supplementation reduces high-glucose and ketone-induced adhesion of monocytes to endothelial cells by inhibiting ROS","volume":"391","author":"Jain","year":"2014","journal-title":"Mol. Cell Biochem."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"625434","DOI":"10.1155\/2012\/625434","article-title":"Hypogonadism in the aging male diagnosis, potential benefits, and risks of testosterone replacement therapy","volume":"2012","author":"Surampudi","year":"2012","journal-title":"Int. J. Endocrinol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1038\/nrurol.2010.230","article-title":"Contemporary concepts in the evaluation and management of male infertility","volume":"8","author":"Hwang","year":"2011","journal-title":"Nat. Rev. Urol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1093\/biolre\/ioy059","article-title":"Leydig cells: Formation, function, and regulation","volume":"99","author":"Zirkin","year":"2018","journal-title":"Biol. Reprod."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"4629","DOI":"10.1210\/en.2014-1688","article-title":"Testosterone treatment is a potent tumor promoter for the rat prostate","volume":"155","author":"Bosland","year":"2014","journal-title":"Endocrinology"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2882","DOI":"10.1210\/en.2002-0090","article-title":"Reactive oxygen disrupts mitochondria in MA-10 tumor Leydig cells and inhibits steroidogenic acute regulatory (StAR) protein and steroidogenesis","volume":"144","author":"Diemer","year":"2003","journal-title":"Endocrinology"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1080\/15376516.2017.1354413","article-title":"T-2 toxin induces apoptosis via the Bax-dependent caspase-3 activation in mouse primary Leydig cells","volume":"28","author":"Zhang","year":"2018","journal-title":"Toxicol. Mech. Methods"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"240","DOI":"10.2164\/jandrol.05075","article-title":"Aging and the brown Norway rat leydig cell antioxidant defense system","volume":"27","author":"Luo","year":"2006","journal-title":"J. Androl."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1361","DOI":"10.1016\/S0531-5565(01)00118-8","article-title":"Age-related increase in mitochondrial superoxide generation in the testosterone-producing cells of Brown Norway rat testes: Relationship to reduced steroidogenic function?","volume":"36","author":"Chen","year":"2001","journal-title":"Exp. Gerontol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1290","DOI":"10.1016\/S0891-5849(02)01040-7","article-title":"Redox analysis of human plasma allows separation of pro-oxidant events of aging from decline in antioxidant defenses","volume":"33","author":"Jones","year":"2002","journal-title":"Free Radic. Biol. Med."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1023\/A:1015643107449","article-title":"Alterations in oxidative stress scavenger system in aging rat brain and lymphocytes","volume":"3","author":"Sandhu","year":"2002","journal-title":"Biogerontology"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1002\/jnr.10217","article-title":"Down-regulation of gamma-glutamylcysteine synthetase regulatory subunit gene expression in rat brain tissue during aging","volume":"68","author":"Liu","year":"2002","journal-title":"J. Neurosci. Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1161\/STROKEAHA.114.008022","article-title":"Dietary cysteine and other amino acids and stroke incidence in women","volume":"46","author":"Larsson","year":"2015","journal-title":"Stroke"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"223","DOI":"10.4103\/1463-1741.137057","article-title":"Comparison of the effects of N-acetyl-cysteine and ginseng in prevention of noise induced hearing loss in male textile workers","volume":"16","author":"Doosti","year":"2014","journal-title":"Noise Health"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1007\/s10741-013-9379-6","article-title":"Prevention of diabetes-induced cardiovascular complications upon treatment with antioxidants","volume":"19","author":"Xu","year":"2014","journal-title":"Heart Fail. Rev."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1002\/ijc.1620","article-title":"Removal of acetaldehyde from saliva by a slow-release buccal tablet of L-cysteine","volume":"97","author":"Salaspuro","year":"2002","journal-title":"Int. J. Cancer"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"5053","DOI":"10.1007\/s12010-023-04457-2","article-title":"Acetylated Oligopeptide and N-acetyl cysteine Protected Against Oxidative Stress, Inflammation, Testicular-Blood Barrier Damage, and Testicular Cell Death in Iron-Overload Rat Model","volume":"195","author":"Ezzat","year":"2023","journal-title":"Appl. Biochem. Biotechnol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1016\/j.abb.2017.07.016","article-title":"l-Cysteine supplementation increases insulin sensitivity mediated by upregulation of GSH and adiponectin in high glucose treated 3T3-L1 adipocytes","volume":"630","author":"Achari","year":"2017","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1080\/07315724.2020.1850371","article-title":"l-Cysteine Stimulates the Effect of Vitamin D on Inhibition of Oxidative Stress, IL-8, and MCP-1 Secretion in High Glucose Treated Monocytes","volume":"40","author":"Jain","year":"2021","journal-title":"J. Am. Coll. Nutr."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2246","DOI":"10.1016\/j.ajpath.2019.07.010","article-title":"N-Acetylcysteine Resolves Placental Inflammatory-Vasculopathic Changes in Mice Consuming a High-Fat Diet","volume":"189","author":"Williams","year":"2019","journal-title":"Am. J. Pathol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.theriogenology.2020.11.005","article-title":"Influence of N-acetylcysteine on steroidogenesis and gene expression in porcine placental trophoblast cells","volume":"161","author":"Ding","year":"2021","journal-title":"Theriogenology"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"882","DOI":"10.5534\/wjmh.220100","article-title":"Effect of N-acetyl-L-cysteine on Testicular Tissue in Busulfan-Induced Dysfunction in the Male Reproductive System","volume":"41","author":"Kim","year":"2023","journal-title":"World J. Mens. Health"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1002\/mrd.22354","article-title":"Alpha-lipoic acid and N-acetylcysteine protects intensive swimming exercise-mediated germ-cell depletion, pro-oxidant generation, and alteration of steroidogenesis in rat testis","volume":"81","author":"Jana","year":"2014","journal-title":"Mol. Reprod. Dev."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Redouane, S., Harmak, H., Elkarhat, Z., Charoute, H., Malki, A., Barakat, A., and Rouba, H. (2023). Exploring the impact of CYP11A1\u2019s missense SNPs on the interaction between CYP11A1 and cholesterol: A comprehensive structural analysis and MD simulation study. Comput. Biol. Chem., 106.","DOI":"10.1016\/j.compbiolchem.2023.107937"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1002\/j.1939-4640.1994.tb00498.x","article-title":"Age-related decreased Leydig cell testosterone production in the brown Norway rat","volume":"15","author":"Chen","year":"1994","journal-title":"J. Androl."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"2773","DOI":"10.1210\/endo.133.6.8243304","article-title":"Reproductive aging in the male brown-Norway rat: A model for the human","volume":"133","author":"Wang","year":"1993","journal-title":"Endocrinology"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1002\/j.1939-4640.2002.tb02251.x","article-title":"Cholesterol transport, peripheral benzodiazepine receptor, and steroidogenesis in aging Leydig cells","volume":"23","author":"Culty","year":"2002","journal-title":"J. Androl."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"820","DOI":"10.1016\/j.jsxm.2018.04.641","article-title":"Testosterone and Cardiovascular Risk: Meta-Analysis of Interventional Studies","volume":"15","author":"Corona","year":"2018","journal-title":"J. Sex. Med."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1016\/j.jsbmb.2016.11.015","article-title":"A time-course regulatory and kinetic expression study of steroid metabolizing enzymes by calcitriol in primary cultured human placental cells","volume":"167","author":"Halhali","year":"2017","journal-title":"J. Steroid Biochem. Mol. Biol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1039\/C8TX00263K","article-title":"Di-(2-ethylhexyl) phthalate (DEHP) inhibits steroidogenesis and induces mitochondria-ROS mediated apoptosis in rat ovarian granulosa cells","volume":"8","author":"Tripathi","year":"2019","journal-title":"Toxicol. Res."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"382","DOI":"10.1093\/toxsci\/kft198","article-title":"Endocrine disruptors differentially target ATP-binding cassette transporters in the blood-testis barrier and affect Leydig cell testosterone secretion in vitro","volume":"136","author":"Dankers","year":"2013","journal-title":"Toxicol. Sci."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.anireprosci.2015.01.007","article-title":"Toxic effects of cadmium on testis of birds and mammals: A review","volume":"155","author":"Marettova","year":"2015","journal-title":"Anim. Reprod. Sci."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Shen, Y., You, Y., Zhu, K., Fang, C., Yu, X., and Chang, D. (2022). Bibliometric and visual analysis of blood-testis barrier research. Front. Pharmacol., 13.","DOI":"10.3389\/fphar.2022.969257"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"316","DOI":"10.1093\/rpd\/ncv270","article-title":"Injury to the blood-testis barrier after low-dose-rate chronic radiation exposure in mice","volume":"167","author":"Son","year":"2015","journal-title":"Radiat. Prot. Dosim."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Lu, Y., Liu, M., Tursi, N.J., Yan, B., Cao, X., Che, Q., Yang, N., and Dong, X. (2021). Uropathogenic Escherichia coli Infection Compromises the Blood-Testis Barrier by Disturbing mTORC1-mTORC2 Balance. Front. Immunol., 12.","DOI":"10.3389\/fimmu.2021.582858"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"8487","DOI":"10.2147\/IJN.S170723","article-title":"Potential adverse effects of nanoparticles on the reproductive system","volume":"13","author":"Wang","year":"2018","journal-title":"Int. J. Nanomed."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Paul, C., and Robaire, B. (2013). Impaired function of the blood-testis barrier during aging is preceded by a decline in cell adhesion proteins and GTPases. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0084354"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"e1009313","DOI":"10.1080\/21565562.2015.1009313","article-title":"Antioxidant protects blood-testis barrier against synchrotron radiation X-ray-induced disruption","volume":"5","author":"Zhang","year":"2015","journal-title":"Spermatogenesis"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"564","DOI":"10.1210\/er.2014-1101","article-title":"The Mammalian Blood-Testis Barrier: Its Biology and Regulation","volume":"36","author":"Mruk","year":"2015","journal-title":"Endocr. Rev."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1186\/1742-2094-11-107","article-title":"Adjudin protects against cerebral ischemia reperfusion injury by inhibition of neuroinflammation and blood-brain barrier disruption","volume":"11","author":"Liu","year":"2014","journal-title":"J. Neuroinflamm."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"6164","DOI":"10.4049\/jimmunol.171.11.6164","article-title":"Proinflammatory cytokines disrupt epithelial barrier function by apoptosis-independent mechanisms","volume":"171","author":"Bruewer","year":"2003","journal-title":"J. Immunol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"864","DOI":"10.1016\/j.bbamem.2008.08.027","article-title":"Cytokine regulation of tight junctions","volume":"1788","author":"Capaldo","year":"2009","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"2153","DOI":"10.1002\/ar.25165","article-title":"The abundance and localization of claudin-1 and -5 in the adult tomcats (Felis catus) testis, tubules rectus, rete testis, efferent ductules, and epididymis","volume":"306","author":"Liman","year":"2023","journal-title":"Anat. Rec."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/j.intimp.2014.08.002","article-title":"THP-1 cell line: An in vitro cell model for immune modulation approach","volume":"23","author":"Chanput","year":"2014","journal-title":"Int. Immunopharmacol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/j.talanta.2018.11.066","article-title":"A series of BODIPY-based probes for the detection of cysteine and homocysteine in living cells","volume":"195","author":"Wang","year":"2019","journal-title":"Talanta"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Clemente Plaza, N., Reig Garcia-Galbis, M., and Martinez-Espinosa, R.M. (2018). Effects of the Usage of l-Cysteine (l-Cys) on Human Health. Molecules, 23.","DOI":"10.3390\/molecules23030575"}],"container-title":["Biomolecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2218-273X\/14\/9\/1171\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T15:58:49Z","timestamp":1760111929000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2218-273X\/14\/9\/1171"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,9,18]]},"references-count":70,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2024,9]]}},"alternative-id":["biom14091171"],"URL":"https:\/\/doi.org\/10.3390\/biom14091171","relation":{},"ISSN":["2218-273X"],"issn-type":[{"value":"2218-273X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,9,18]]}}}