{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T23:53:47Z","timestamp":1790985227960,"version":"4.1.0"},"reference-count":242,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,6,8]],"date-time":"2026-06-08T00:00:00Z","timestamp":1780876800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T00:00:00Z","timestamp":1790899200000},"content-version":"vor","delay-in-days":116,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Biol Res"],"abstract":"<jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>Ovarian theca cells constitute essential components of the follicular microenvironment and play central roles in follicular development, steroidogenesis, and endocrine regulation. Despite their significance, the developmental origins, differentiation processes, and functional dynamics of theca cells remain incompletely defined, particularly in humans. This review provides an updated synthesis of current knowledge on the ontogeny, molecular signaling pathways, and intercellular interactions of theca cells. It also presents recent findings on the role of theca stem or progenitor cells and their relevance to reproductive disorders, including polycystic ovary syndrome (PCOS), hyperthecosis, and ovarian insufficiency. A comprehensive literature search was conducted using PubMed, Scopus, and Web of Science through May 2025. Search terms included \u201ctheca cells,\u201d \u201ctheca progenitors,\u201d \u201cfollicular development,\u201d \u201covarian differentiation,\u201d \u201csteroidogenesis,\u201d and \u201creproductive disorders.\u201d Original research articles and reviews providing mechanistic insights were included. Current evidence indicates that several signaling pathways, including IGF, SCF, FGF, members of the TGF-\u03b2 superfamily, GDF9, BMPs, and Hedgehog proteins, are involved in the recruitment, proliferation, and differentiation of theca cells. Bidirectional communication among oocytes, granulosa cells, and theca cells remains essential for folliculogenesis. Increasing evidence links theca cell dysfunction to the pathophysiology of PCOS, primary ovarian insufficiency, and reproductive aging. The identification of theca stem cells (TSCs) and their proposed roles in ovarian regeneration represents an important conceptual advance. A deeper understanding of theca cell biology may guide the development of targeted strategies for infertility and ovarian dysfunction. In contrast, the integration of TSC biology offers new directions for reproductive medicine and reproductive health.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s40659-026-00680-3","type":"journal-article","created":{"date-parts":[[2026,6,8]],"date-time":"2026-06-08T07:26:03Z","timestamp":1780903563000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Developmental and functional landscape of theca cells in ovarian biology"],"prefix":"10.1186","volume":"59","author":[{"given":"Mahnaz","family":"Haddadi","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samane","family":"Adib","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christiani A.","family":"Amorim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5891-3094","authenticated-orcid":false,"given":"Azam","family":"Dalman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6592-8243","authenticated-orcid":false,"given":"Mojtaba Rezazadeh","family":"Valojerdi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,6,8]]},"reference":[{"issue":"4","key":"680_CR1","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1530\/REP-10-0094","volume":"140","author":"J Young","year":"2010","unstructured":"Young J, McNeilly AS. Theca: the forgotten cell of the ovarian follicle. Reproduction. 2010;140(4):489.","journal-title":"Reproduction"},{"issue":"2","key":"680_CR2","doi-asserted-by":"publisher","first-page":"1128","DOI":"10.1111\/jcmm.16179","volume":"25","author":"HY Zhang","year":"2021","unstructured":"Zhang HY, Zhu FF, Zhu YJ, Hu YJ, Chen X. Effects of IL-18 on the proliferation and steroidogenesis of bovine theca cells: possible roles in the pathogenesis of polycystic ovary syndrome. J Cell Mol Med. 2021;25(2):1128\u201339.","journal-title":"J Cell Mol Med"},{"issue":"12","key":"680_CR3","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21124565","volume":"21","author":"E Gershon","year":"2020","unstructured":"Gershon E, Dekel N. Newly identified regulators of ovarian folliculogenesis and ovulation. Int J Mol Sci. 2020;21(12):4565.","journal-title":"Int J Mol Sci"},{"issue":"14","key":"680_CR4","doi-asserted-by":"publisher","first-page":"1711","DOI":"10.1097\/CM9.0000000000000850","volume":"133","author":"T Liu","year":"2020","unstructured":"Liu T, Qin Q-Y, Qu J-X, Wang H-Y, Yan J. Where are the theca cells from: the mechanism of theca cells derivation and differentiation. Chin Med J. 2020;133(14):1711\u20138.","journal-title":"Chin Med J"},{"issue":"1","key":"680_CR5","doi-asserted-by":"publisher","first-page":"174","DOI":"10.3390\/cells12010174","volume":"12","author":"M Jozkowiak","year":"2022","unstructured":"Jozkowiak M, Piotrowska-Kempisty H, Kobylarek D, Gorska N, Mozdziak P, Kempisty B, et al. Endocrine disrupting chemicals in polycystic ovary syndrome: the relevant role of the theca and granulosa cells in the pathogenesis of the ovarian dysfunction. Cells. 2022;12(1):174.","journal-title":"Cells"},{"issue":"2","key":"680_CR6","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1016\/j.fertnstert.2020.08.005","volume":"115","author":"C Zhu","year":"2021","unstructured":"Zhu C, Luo W, Li Z, Zhang X, Hu J, Zhao S, et al. New theca-cell marker insulin-like factor 3 is associated with premature ovarian insufficiency. Fertil Steril. 2021;115(2):455\u201362.","journal-title":"Fertil Steril"},{"issue":"2","key":"680_CR7","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1093\/stmcls\/sxac070","volume":"41","author":"M Yoshihara","year":"2023","unstructured":"Yoshihara M, Wagner M, Damdimopoulos A, Zhao C, Petropoulos S, Katayama S, et al. The continued absence of functional germline stem cells in adult ovaries. Stem Cells. 2023;41(2):105\u201310.","journal-title":"Stem Cells"},{"issue":"9","key":"680_CR8","doi-asserted-by":"publisher","first-page":"1150","DOI":"10.1038\/s41559-020-1262-y","volume":"4","author":"TC Wanger","year":"2020","unstructured":"Wanger TC, DeClerck F, Garibaldi LA, Ghazoul J, Kleijn D, Klein A-M, et al. Integrating agroecological production in a robust post-2020 global biodiversity framework. Nat Ecol Evolution. 2020;4(9):1150\u20132.","journal-title":"Nat Ecol Evolution"},{"key":"680_CR9","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1007\/978-981-19-8848-6_2","volume-title":"The life cycle of the ovary. Ovarian aging","author":"L Ma","year":"2023","unstructured":"Ma L, Shen W, Zhang J, Ma L, Shen W, Shen W, et al. The life cycle of the ovary. Ovarian aging. New York: Springer; 2023. p. 7\u201333."},{"key":"680_CR10","doi-asserted-by":"publisher","DOI":"10.3389\/fendo.2019.00832","volume":"10","author":"MA Abedal-Majed","year":"2019","unstructured":"Abedal-Majed MA, Romereim S, Davis JS, Cupp AS. Perturbations in lineage specification of granulosa and theca cells may alter corpus luteum formation and function. Front Endocrinol. 2019;10:832.","journal-title":"Front Endocrinol"},{"issue":"4","key":"680_CR11","first-page":"1158","volume":"79","author":"C Gilling-Smith","year":"1994","unstructured":"Gilling-Smith C, Willis DS, Beard RW, Franks S. Hypersecretion of androstenedione by isolated thecal cells from polycystic ovaries. J Clin Endocrinol Metab. 1994;79(4):1158\u201365.","journal-title":"J Clin Endocrinol Metab"},{"key":"680_CR12","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1007\/978-981-16-9016-7_9","volume-title":"Emerging concepts in endocrine structure and functions","author":"EA Al-Suhaimi","year":"2022","unstructured":"Al-Suhaimi EA, Khan FA, Homeida A. Regulation of male and female reproductive functions. In: Emerging concepts in endocrine structure and functions. New York: Springer; 2022. p. 287\u2013347."},{"issue":"4","key":"680_CR13","doi-asserted-by":"publisher","first-page":"920","DOI":"10.1210\/en.2016-1538","volume":"158","author":"M Laird","year":"2017","unstructured":"Laird M, Thomson K, Fenwick M, Mora J, Franks S, Hardy K. Androgen stimulates growth of mouse preantral follicles in vitro: interaction with follicle-stimulating hormone and with growth factors of the TGF \u03b2 superfamily. Endocrinology. 2017;158(4):920\u201335.","journal-title":"Endocrinology"},{"issue":"4","key":"680_CR14","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1002\/mrd.23683","volume":"90","author":"A Guzm\u00e1n","year":"2023","unstructured":"Guzm\u00e1n A, Hern\u00e1ndez\u2010Coronado CG, Guti\u00e9rrez CG, Rosales\u2010Torres AM. The vascular endothelial growth factor (VEGF) system as a key regulator of ovarian follicle angiogenesis and growth. Mol Reprod Dev. 2023;90(4):201\u201317.","journal-title":"Mol Reprod Dev"},{"issue":"1","key":"680_CR15","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1210\/er.2017-00164","volume":"39","author":"JS Richards","year":"2018","unstructured":"Richards JS, Ren YA, Candelaria N, Adams JE, Rajkovic A. Ovarian follicular theca cell recruitment, differentiation, and impact on fertility: 2017 update. Endocr Rev. 2018;39(1):1\u201320.","journal-title":"Endocr Rev"},{"issue":"1","key":"680_CR16","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-25837-3","volume":"8","author":"Y Ono","year":"2018","unstructured":"Ono Y, Nagai M, Yoshino O, Koga K, Nawaz A, Hatta H, et al. CD11c+ M1-like macrophages (M\u03a6s) but not CD206+ M2-like M\u03a6 are involved in folliculogenesis in mice ovary. Sci Rep. 2018;8(1):8171.","journal-title":"Sci Rep"},{"issue":"1","key":"680_CR17","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-025-65390-y","volume":"16","author":"A Biswas","year":"2025","unstructured":"Biswas A, Lou Y, Ng BH, Tomida K, Darpe S, Leong KW, et al. Surface mechanics and compressive stress impact mammalian follicle development. Nat Commun. 2025;16(1):9578.","journal-title":"Nat Commun"},{"key":"680_CR18","doi-asserted-by":"publisher","DOI":"10.3389\/fcell.2022.1038107","volume":"10","author":"A Biswas","year":"2022","unstructured":"Biswas A, Ng BH, Prabhakaran VSO, Chan CJ. Squeezing the eggs to grow: the mechanobiology of mammalian folliculogenesis. Front Cell Dev Biol. 2022;10:1038107.","journal-title":"Front Cell Dev Biol"},{"issue":"1\u20132","key":"680_CR19","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1016\/S0303-7207(01)00469-5","volume":"179","author":"S Hillier","year":"2001","unstructured":"Hillier S. Gonadotropic control of ovarian follicular growth and development. Mol Cell Endocrinol. 2001;179(1\u20132):39\u201346.","journal-title":"Mol Cell Endocrinol"},{"issue":"4","key":"680_CR20","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1007\/BF00324744","volume":"85","author":"J Priedkalns","year":"1968","unstructured":"Priedkalns J, Weber AF, Zemjanis R. Qualitative and quantitative morphological studies of the cells of the membrana granulosa, theca interna and corpus luteum of the bovine ovary. Z Zellforsch Mikrosk Anat. 1968;85(4):501\u201320.","journal-title":"Z Zellforsch Mikrosk Anat"},{"issue":"3","key":"680_CR21","doi-asserted-by":"publisher","first-page":"787","DOI":"10.1258\/0007142001903364","volume":"56","author":"HM Fraser","year":"2000","unstructured":"Fraser HM, Lunn SF. Angiogenesis and its control in the female reproductive system. Br Med Bull. 2000;56(3):787\u201397.","journal-title":"Br Med Bull"},{"issue":"5","key":"680_CR22","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1007\/s00418-006-0194-1","volume":"126","author":"CB Berkholtz","year":"2006","unstructured":"Berkholtz CB, Lai BE, Woodruff TK, Shea LD. Distribution of extracellular matrix proteins type I collagen, type IV collagen, fibronectin, and laminin in mouse folliculogenesis. Histochem Cell Biol. 2006;126(5):583\u201392.","journal-title":"Histochem Cell Biol"},{"issue":"12","key":"680_CR23","doi-asserted-by":"publisher","first-page":"786","DOI":"10.1038\/nrm3904","volume":"15","author":"C Bonnans","year":"2014","unstructured":"Bonnans C, Chou J, Werb Z. Remodelling the extracellular matrix in development and disease. Nat Rev Mol Cell Biol. 2014;15(12):786\u2013801.","journal-title":"Nat Rev Mol Cell Biol"},{"issue":"7","key":"680_CR24","doi-asserted-by":"publisher","first-page":"3403","DOI":"10.3390\/ijms22073403","volume":"22","author":"P Carey","year":"2021","unstructured":"Carey P, Low E, Harper E, Stack MS. Metalloproteinases in ovarian cancer. Int J Mol Sci. 2021;22(7):3403.","journal-title":"Int J Mol Sci"},{"issue":"1","key":"680_CR25","doi-asserted-by":"publisher","first-page":"54","DOI":"10.3390\/ijms20010054","volume":"20","author":"K Sterzynska","year":"2018","unstructured":"Sterzynska K, Klejewski A, Wojtowicz K, Swierczewska M, Nowacka M, Kazmierczak D, et al. Mutual expression of ALDH1A1, LOX, and collagens in ovarian cancer cell lines as combined CSCs-and ECM-related models of drug resistance development. Int J Mol Sci. 2018;20(1):54.","journal-title":"Int J Mol Sci"},{"issue":"2","key":"680_CR26","first-page":"285","volume":"17","author":"MS \u00dcnal","year":"2024","unstructured":"\u00dcnal MS, Tan S, Se\u00e7me M. Changes in structure during the corpus luteum\u2019s formation. Pamukkale Med J. 2024;17(2):285\u2013301.","journal-title":"Pamukkale Med J"},{"issue":"19","key":"680_CR27","doi-asserted-by":"publisher","first-page":"4947","DOI":"10.3390\/ijms20194947","volume":"20","author":"M Paolillo","year":"2019","unstructured":"Paolillo M, Schinelli S. Extracellular matrix alterations in metastatic processes. Int J Mol Sci. 2019;20(19):4947.","journal-title":"Int J Mol Sci"},{"issue":"12","key":"680_CR28","doi-asserted-by":"publisher","first-page":"2793","DOI":"10.1093\/humrep\/deaa246","volume":"35","author":"P Asiabi","year":"2020","unstructured":"Asiabi P, Dolmans M-M, Ambroise J, Camboni A, Amorim C. In vitro differentiation of theca cells from ovarian cells isolated from postmenopausal women. Hum Reprod. 2020;35(12):2793\u2013807.","journal-title":"Hum Reprod"},{"issue":"7","key":"680_CR29","doi-asserted-by":"publisher","first-page":"1344","DOI":"10.1016\/j.biocel.2005.01.016","volume":"37","author":"DA Magoffin","year":"2005","unstructured":"Magoffin DA. Ovarian theca cell. Int J Biochem Cell Biol. 2005;37(7):1344\u20139.","journal-title":"Int J Biochem Cell Biol"},{"key":"680_CR30","doi-asserted-by":"publisher","first-page":"106","DOI":"10.3389\/fendo.2016.00106","volume":"7","author":"A Midzak","year":"2016","unstructured":"Midzak A, Papadopoulos V. Adrenal mitochondria and steroidogenesis: from individual proteins to functional protein assemblies. Front Endocrinol. 2016;7:106.","journal-title":"Front Endocrinol"},{"issue":"4","key":"680_CR31","doi-asserted-by":"publisher","first-page":"e70006","DOI":"10.1002\/qub2.70006","volume":"13","author":"S Mao","year":"2025","unstructured":"Mao S, Zhang C, Chen R, Tang S, Fan X, Hu J. Cell lineage tracing: methods, applications, and challenges. Quant Biol. 2025;13(4):e70006.","journal-title":"Quant Biol"},{"issue":"10","key":"680_CR32","doi-asserted-by":"publisher","first-page":"a035618","DOI":"10.1101\/cshperspect.a035618","volume":"12","author":"RN Das","year":"2020","unstructured":"Das RN, Yaniv K. Discovering new progenitor cell populations through lineage tracing and in vivo imaging. Cold Spring Harb Perspect Biol. 2020;12(10):a035618.","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"680_CR33","first-page":"211","volume-title":"Germline development: methods and protocols","author":"C Liu","year":"2011","unstructured":"Liu C, Paczkowski M, Othman M, Yao HH-C. Investigating the origins of somatic cell populations in the perinatal mouse ovaries using genetic lineage tracing and immunohistochemistry. Germline development: methods and protocols. New York: Springer; 2011. pp. 211\u201321."},{"key":"680_CR34","first-page":"19","volume-title":"Engineering in translational medicine","author":"RK Singh","year":"2013","unstructured":"Singh RK, Gaikwad SM, Chatterjee S, Ray P. Stem cells: the holy grail of regenerative medicine. Engineering in translational medicine. New York: Springer; 2013. pp. 19\u201369."},{"issue":"7","key":"680_CR35","doi-asserted-by":"publisher","first-page":"656","DOI":"10.1007\/s11626-014-9750-5","volume":"50","author":"M Nagyova","year":"2014","unstructured":"Nagyova M, Slovinska L, Blasko J, Grulova I, Kuricova M, Cigankova V, et al. A comparative study of PKH67, DiI, and BrdU labeling techniques for tracing rat mesenchymal stem cells. Vitro Cell Dev Biol. 2014;50(7):656\u201363.","journal-title":"Vitro Cell Dev Biol"},{"issue":"5","key":"680_CR36","doi-asserted-by":"publisher","first-page":"1234","DOI":"10.1093\/biolre\/ioab163","volume":"105","author":"S Li","year":"2021","unstructured":"Li S, Chen L-N, Zhu H-J, Feng X, Xie F-Y, Luo S-M, et al. Single-cell RNA sequencing analysis of mouse follicular somatic cells. Biol Reprod. 2021;105(5):1234\u201345.","journal-title":"Biol Reprod"},{"issue":"1","key":"680_CR37","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1038\/s41421-022-00492-1","volume":"8","author":"J Zhou","year":"2022","unstructured":"Zhou J, Jiang X, Wu H, Zhang L, Chen M, Chen M, et al. Dissecting the fate of Foxl2-expressing cells in fetal ovary using lineage tracing and single-cell transcriptomics. Cell Discov. 2022;8(1):139.","journal-title":"Cell Discov"},{"issue":"12","key":"680_CR38","doi-asserted-by":"publisher","first-page":"e3001025","DOI":"10.1371\/journal.pbio.3001025","volume":"18","author":"J-J Wang","year":"2020","unstructured":"Wang J-J, Ge W, Zhai Q-Y, Liu J-C, Sun X-W, Liu W-X, et al. Single-cell transcriptome landscape of ovarian cells during primordial follicle assembly in mice. PLoS Biol. 2020;18(12):e3001025.","journal-title":"PLoS Biol"},{"issue":"15","key":"680_CR39","doi-asserted-by":"publisher","first-page":"1248","DOI":"10.3390\/cells13151248","volume":"13","author":"H Vlieghe","year":"2024","unstructured":"Vlieghe H, Sousa MJ, Charif D, Amorim CA. Unveiling the differentiation potential of ovarian theca interna cells from multipotent stem cell-like cells. Cells. 2024;13(15):1248.","journal-title":"Cells"},{"issue":"1","key":"680_CR40","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/ncomms7934","volume":"6","author":"C Liu","year":"2015","unstructured":"Liu C, Peng J, Matzuk MM, Yao HH-C. Lineage specification of ovarian theca cells requires multicellular interactions via oocyte and granulosa cells. Nat Commun. 2015;6(1):1\u201311.","journal-title":"Nat Commun"},{"issue":"30","key":"680_CR41","doi-asserted-by":"publisher","first-page":"12389","DOI":"10.1073\/pnas.0703787104","volume":"104","author":"A Honda","year":"2007","unstructured":"Honda A, Hirose M, Hara K, Matoba S, Inoue K, Miki H, et al. Isolation, characterization, and in vitro and in vivo differentiation of putative thecal stem cells. Proc Natl Acad Sci. 2007;104(30):12389\u201394.","journal-title":"Proc Natl Acad Sci"},{"issue":"8","key":"680_CR42","doi-asserted-by":"publisher","first-page":"3789","DOI":"10.1210\/en.2006-0394","volume":"147","author":"WL Dees","year":"2006","unstructured":"Dees WL, Hiney J, McArthur N, Johnson G, Dissen G, Ojeda S. Origin and ontogeny of mammalian ovarian neurons. Endocrinology. 2006;147(8):3789\u201396.","journal-title":"Endocrinology"},{"issue":"12","key":"680_CR43","doi-asserted-by":"publisher","first-page":"5570","DOI":"10.1073\/pnas.1814930116","volume":"116","author":"J McKey","year":"2019","unstructured":"McKey J, Bunce C, Batchvarov IS, Ornitz DM, Capel B. Neural crest-derived neurons invade the ovary but not the testis during mouse gonad development. Proc Natl Acad Sci USA. 2019;116(12):5570\u20135.","journal-title":"Proc Natl Acad Sci USA"},{"issue":"1","key":"680_CR44","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1210\/er.2014-1079","volume":"36","author":"K Hummitzsch","year":"2015","unstructured":"Hummitzsch K, Anderson RA, Wilhelm D, Wu J, Telfer EE, Russell DL, et al. Stem cells, progenitor cells, and lineage decisions in the ovary. Endocr Rev. 2015;36(1):65\u201391.","journal-title":"Endocr Rev"},{"issue":"5","key":"680_CR45","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1210\/er.2018-00010","volume":"39","author":"E Rotgers","year":"2018","unstructured":"Rotgers E, J\u00f8rgensen A, Yao HH-C. At the crossroads of fate\u2014somatic cell lineage specification in the fetal gonad. Endocr Rev. 2018;39(5):739\u201359.","journal-title":"Endocr Rev"},{"issue":"1","key":"680_CR46","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1016\/j.matbio.2011.10.002","volume":"31","author":"N Hatzirodos","year":"2012","unstructured":"Hatzirodos N, Nigro J, Irving-Rodgers HF, Vashi AV, Hummitzsch K, Caterson B, et al. Glycomic analyses of ovarian follicles during development and atresia. Matrix Biol. 2012;31(1):45\u201356.","journal-title":"Matrix Biol"},{"key":"680_CR47","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1016\/j.mce.2016.09.029","volume":"439","author":"SM Romereim","year":"2017","unstructured":"Romereim SM, Summers AF, Pohlmeier WE, Zhang P, Hou X, Talbott HA, et al. Gene expression profiling of bovine ovarian follicular and luteal cells provides insight into cellular identities and functions. Mol Cell Endocrinol. 2017;439:379\u201394.","journal-title":"Mol Cell Endocrinol"},{"issue":"1","key":"680_CR48","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-022-04384-8","volume":"6","author":"NL Guahmich","year":"2023","unstructured":"Guahmich NL, Man L, Wang J, Arazi L, Kallinos E, Topper-Kroog A, et al. Human theca arises from ovarian stroma and is comprised of three discrete subtypes. Commun Biol. 2023;6(1):7.","journal-title":"Commun Biol"},{"issue":"7","key":"680_CR49","doi-asserted-by":"publisher","first-page":"755","DOI":"10.1101\/gad.13.7.755","volume":"13","author":"A Swain","year":"1999","unstructured":"Swain A, Lovell-Badge R. Mammalian sex determination: a molecular drama. Genes Dev. 1999;13(7):755\u201367.","journal-title":"Genes Dev"},{"issue":"3","key":"680_CR50","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1016\/j.tvjl.2013.04.011","volume":"197","author":"Y-M Lee","year":"2013","unstructured":"Lee Y-M, Kumar BM, Lee J-H, Lee W-J, Kim T-H, Lee S-L, et al. Characterisation and differentiation of porcine ovarian theca-derived multipotent stem cells. Vet J. 2013;197(3):761\u20138.","journal-title":"Vet J"},{"key":"680_CR51","doi-asserted-by":"publisher","first-page":"378","DOI":"10.1016\/j.rvsc.2017.06.021","volume":"114","author":"S Adib","year":"2017","unstructured":"Adib S, Valojerdi MR. Molecular assessment, characterization, and differentiation of theca stem cells imply the presence of mesenchymal and pluripotent stem cells in sheep ovarian theca layer. Res Vet Sci. 2017;114:378\u201387.","journal-title":"Res Vet Sci"},{"issue":"4","key":"680_CR52","volume":"20","author":"A Dalman","year":"2019","unstructured":"Dalman A, Totonchi M, Valojerdi MR. Human ovarian theca-derived multipotent stem cells have the potential to differentiate into oocyte-like cells in vitro. Cell J. 2019;20(4):527.","journal-title":"Cell J"},{"issue":"7","key":"680_CR53","doi-asserted-by":"publisher","first-page":"1611","DOI":"10.1007\/s10815-023-02799-8","volume":"40","author":"A Dalman","year":"2023","unstructured":"Dalman A, Adib S, Amorim CA, Pirjani R, Totonchi M, Valojerdi MR. Co-culture of human cryopreserved fragmented ovarian tissue with theca progenitor cells derived from theca stem cells. J Assist Reprod Genet. 2023;40(7):1611\u201322.","journal-title":"J Assist Reprod Genet"},{"issue":"1","key":"680_CR54","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1186\/s13048-023-01315-x","volume":"17","author":"SN Mirbahari","year":"2024","unstructured":"Mirbahari SN, Amorim CA, Hassani F, Totonchi M, Haddadi M, Valojerdi MR, et al. In-vitro generation of follicle-like structures from human germ cell-like cells derived from theca stem cell combined with ovarian somatic cells. J Ovarian Res. 2024;17(1):2.","journal-title":"J Ovarian Res"},{"issue":"3","key":"680_CR55","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1262\/jrd.2014-107","volume":"61","author":"N Shiomi-Sugaya","year":"2015","unstructured":"Shiomi-Sugaya N, Komatsu K, Wang J, Yamashita M, Kikkawa F, Iwase A. Regulation of secondary follicle growth by theca cells and insulin-like growth factor 1. J Reprod Dev. 2015;61(3):161\u20138.","journal-title":"J Reprod Dev"},{"issue":"2","key":"680_CR56","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1095\/biolreprod64.2.451","volume":"64","author":"CT Huang","year":"2001","unstructured":"Huang CT, Weitsman SR, Dykes BN, Magoffin DA. Stem cell factor and insulin-like growth factor-I stimulate luteinizing hormone-independent differentiation of rat ovarian theca cells. Biol Reprod. 2001;64(2):451\u20136.","journal-title":"Biol Reprod"},{"issue":"2","key":"680_CR57","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1016\/S0015-0282(97)00473-1","volume":"69","author":"AJ Duleba","year":"1998","unstructured":"Duleba AJ, Spaczynski RZ, Olive DL. Insulin and insulin-like growth factor I stimulate the proliferation of human ovarian theca-interstitial cells. Fertil Steril. 1998;69(2):335\u201340.","journal-title":"Fertil Steril"},{"issue":"4","key":"680_CR58","doi-asserted-by":"publisher","first-page":"891","DOI":"10.1095\/biolreprod56.4.891","volume":"56","author":"AJ Duleba","year":"1997","unstructured":"Duleba AJ, Spaczynski RZ, Olive DL, Behrman HR. Effects of insulin and insulin-like growth factors on proliferation of rat ovarian theca-interstitial cells. Biol Reprod. 1997;56(4):891\u20137.","journal-title":"Biol Reprod"},{"issue":"3","key":"680_CR59","doi-asserted-by":"publisher","first-page":"546","DOI":"10.1095\/biolreprod60.3.546","volume":"60","author":"AJ Duleba","year":"1999","unstructured":"Duleba AJ, Spaczynski RZ, Arici A, Carbone R, Behrman HR. Proliferation and differentiation of rat theca-interstitial cells: comparison of effects induced by platelet-derived growth factor and insulin-like growth factor-I. Biol Reprod. 1999;60(3):546\u201350.","journal-title":"Biol Reprod"},{"issue":"1","key":"680_CR60","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1095\/biolreprod54.1.255","volume":"54","author":"L Spicer","year":"1996","unstructured":"Spicer L, Stewart R. Interactions among basic fibroblast growth factor, epidermal growth factor, insulin, and insulin-like growth factor-I (IGF-I) on cell numbers and steroidogenesis of bovine thecal cells: role of IGF-I receptors. Biol Reprod. 1996;54(1):255\u201363.","journal-title":"Biol Reprod"},{"issue":"5","key":"680_CR61","doi-asserted-by":"publisher","first-page":"973","DOI":"10.1016\/S0015-0282(01)01747-2","volume":"75","author":"M Ito","year":"2001","unstructured":"Ito M, Harada T, Tanikawa M, Fujii A, Shiota G, Terakawa N. Hepatocyte growth factor and stem cell factor involvement in paracrine interplays of theca and granulosa cells in the human ovary. Fertil Steril. 2001;75(5):973\u20139.","journal-title":"Fertil Steril"},{"issue":"1-2","key":"680_CR62","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.mce.2004.09.012","volume":"234","author":"PE H\u00f8yer","year":"2005","unstructured":"H\u00f8yer PE, Byskov AG, M\u00f8llg\u00e5rd K. Stem cell factor and c-Kit in human primordial germ cells and fetal ovaries. Mol Cell Endocrinol. 2005;234(1\u20132):1\u201310.","journal-title":"Mol Cell Endocrinol"},{"issue":"6","key":"680_CR63","doi-asserted-by":"publisher","first-page":"647","DOI":"10.1016\/j.acthis.2010.08.004","volume":"113","author":"F Raucci","year":"2011","unstructured":"Raucci F, Di Fiore MM. Localization of c-kit and stem cell factor (SCF) in ovarian follicular epithelium of a lizard, Podarcis s. sicula. Acta Histochem. 2011;113(6):647\u201355.","journal-title":"Acta Histochem"},{"issue":"1-2","key":"680_CR64","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/j.mce.2004.10.006","volume":"229","author":"X Jin","year":"2005","unstructured":"Jin X, Han C-S, Zhang X-S, Yuan J-X, Hu Z-Y, Liu Y-X. Signal transduction of stem cell factor in promoting early follicle development. Mol Cell Endocrinol. 2005;229(1\u20132):3\u201310.","journal-title":"Mol Cell Endocrinol"},{"key":"680_CR65","doi-asserted-by":"publisher","DOI":"10.1016\/j.domaniend.2022.106712","volume":"80","author":"L Sch\u00fctz","year":"2022","unstructured":"Sch\u00fctz L, Hemple A, Morrell B, Schreiber N, Gilliam J, Cortinovis C, et al. Changes in fibroblast growth factor receptors-1c,-2c,-3c, and-4 mRNA in granulosa and theca cells during ovarian follicular growth in dairy cattle. Domest Anim Endocrinol. 2022;80:106712.","journal-title":"Domest Anim Endocrinol"},{"key":"680_CR66","doi-asserted-by":"publisher","first-page":"1250551","DOI":"10.3389\/fvets.2023.1250551","volume":"10","author":"H Zheng","year":"2023","unstructured":"Zheng H, Choi H, Oh D, Kim M, Cai L, Jawad A, et al. Supplementation with fibroblast growth factor 7 during in vitro maturation of porcine cumulus-oocyte complexes improves oocyte maturation and early embryonic development. Front Vet Sci. 2023;10:1250551.","journal-title":"Front Vet Sci"},{"issue":"1","key":"680_CR67","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1016\/j.xfss.2023.10.005","volume":"5","author":"X Guo","year":"2024","unstructured":"Guo X, Zhong Y, Liu Y, Wu R, Huang L, Huang C, et al. Oocyte-derived growth differentiation factor 9 suppresses the expression of CYP17A1 and androgen production in human theca cells. F&S Sci. 2024;5(1):16\u201323.","journal-title":"F&S Sci"},{"issue":"5","key":"680_CR68","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1530\/REP-10-0216","volume":"140","author":"CSL Glister","year":"2010","unstructured":"Glister CSL, Knight PG. Changes in expression of bone morphogenetic proteins (BMPs), their receptors and inhibin co-receptor betaglycan during bovine antral follicle development: inhibin can antagonize the suppressive effect of BMPs on thecal androgen production. Reproduction. 2010;140(5):699\u2013712.","journal-title":"Reproduction"},{"issue":"1","key":"680_CR69","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1016\/0002-9378(90)90864-4","volume":"162","author":"WN Burns","year":"1990","unstructured":"Burns WN, McGill JR, Roy AK, Schenken RS. Expression of the human inhibin \u03b1-subunit gene in preovulatory granulosa-theca cells. Am J Obstet Gynecol. 1990;162(1):273\u20137.","journal-title":"Am J Obstet Gynecol"},{"issue":"17","key":"680_CR70","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22179184","volume":"22","author":"RM Chugh","year":"2021","unstructured":"Chugh RM, Park H-s, Esfandyari S, Elsharoud A, Ulin M, Al-Hendy A. Mesenchymal stem cell-conditioned media regulate steroidogenesis and inhibit androgen secretion in a PCOS cell model via BMP-2. Int J Mol Sci. 2021;22(17):9184.","journal-title":"Int J Mol Sci"},{"issue":"10","key":"680_CR71","doi-asserted-by":"publisher","first-page":"3814","DOI":"10.1210\/endo.141.10.7732","volume":"141","author":"UA Vitt","year":"2000","unstructured":"Vitt UA, McGee EA, Hayashi M, Hsueh AJ. In vivo treatment with GDF-9 stimulates primordial and primary follicle progression and theca cell marker CYP17 in ovaries of immature rats. Endocrinology. 2000;141(10):3814\u201320.","journal-title":"Endocrinology"},{"issue":"5","key":"680_CR72","doi-asserted-by":"publisher","first-page":"1444","DOI":"10.1016\/j.fertnstert.2013.07.200","volume":"100","author":"M-h Chen","year":"2013","unstructured":"Chen M-h, Li T, Ding C-h, Xu Y-w, Guo L-y, Zhou C-q. Growth differential factor-9 inhibits testosterone production in mouse theca interstitial cells. Fertil Steril. 2013;100(5):1444\u201350.","journal-title":"Fertil Steril"},{"issue":"8","key":"680_CR73","doi-asserted-by":"publisher","first-page":"3558","DOI":"10.1210\/en.2005-0311","volume":"146","author":"M Wijgerde","year":"2005","unstructured":"Wijgerde M, Ooms M, Hoogerbrugge JW, Grootegoed JA. Hedgehog signaling in mouse ovary: Indian hedgehog and desert hedgehog from granulosa cells induce target gene expression in developing theca cells. Endocrinology. 2005;146(8):3558\u201366.","journal-title":"Endocrinology"},{"issue":"2","key":"680_CR74","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1530\/REP-08-0317","volume":"138","author":"LJ Spicer","year":"2009","unstructured":"Spicer LJ, Sudo S, Aad PY, Wang LS, Chun S-Y, Ben-Shlomo I, et al. The hedgehog-patched signaling pathway and function in the mammalian ovary: a novel role for hedgehog proteins in stimulating proliferation and steroidogenesis of theca cells. Reproduction. 2009;138(2):329\u201339.","journal-title":"Reproduction"},{"issue":"4","key":"680_CR75","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1530\/REP-20-0471","volume":"161","author":"RG Cowan","year":"2021","unstructured":"Cowan RG, Quirk SM. Cells responding to hedgehog signaling contribute to the theca of ovarian follicles. Reproduction. 2021;161(4):437\u201348.","journal-title":"Reproduction"},{"issue":"1","key":"680_CR76","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1186\/s43043-022-00119-1","volume":"27","author":"P Afradiasbagharani","year":"2022","unstructured":"Afradiasbagharani P, Hosseini E, Allahveisi A, Bazrafkan M. The insulin-like growth factor and its players: their functions, significance, and consequences in all aspects of ovarian physiology. Middle East Fertil Soc J. 2022;27(1):27.","journal-title":"Middle East Fertil Soc J"},{"key":"680_CR77","doi-asserted-by":"publisher","first-page":"777","DOI":"10.3389\/fendo.2019.00777","volume":"10","author":"E Ipsa","year":"2019","unstructured":"Ipsa E, Cruzat VF, Kagize JN, Yovich JL, Keane KN. Growth hormone and insulin-like growth factor action in reproductive tissues. Front Endocrinol. 2019;10:777.","journal-title":"Front Endocrinol"},{"key":"680_CR78","doi-asserted-by":"publisher","first-page":"450","DOI":"10.3389\/fendo.2019.00450","volume":"10","author":"J Devesa","year":"2019","unstructured":"Devesa J, Caicedo D. The role of growth hormone on ovarian functioning and ovarian angiogenesis. Front Endocrinol. 2019;10:450.","journal-title":"Front Endocrinol"},{"key":"680_CR79","doi-asserted-by":"publisher","first-page":"1040503","DOI":"10.3389\/fendo.2022.1040503","volume":"13","author":"C-W Chang","year":"2022","unstructured":"Chang C-W, Sung Y-W, Hsueh Y-W, Chen Y-Y, Ho M, Hsu H-C, et al. Growth hormone in fertility and infertility: mechanisms of action and clinical applications. Front Endocrinol. 2022;13:1040503.","journal-title":"Front Endocrinol"},{"issue":"1\u20132","key":"680_CR80","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1016\/j.anireprosci.2007.04.005","volume":"106","author":"T Shimizu","year":"2008","unstructured":"Shimizu T, Murayama C, Sudo N, Kawashima C, Tetsuka M, Miyamoto A. Involvement of insulin and growth hormone (GH) during follicular development in the bovine ovary. Anim Reprod Sci. 2008;106(1\u20132):143\u201352.","journal-title":"Anim Reprod Sci"},{"key":"680_CR81","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1016\/j.canlet.2017.07.017","volume":"405","author":"MI Figueira","year":"2017","unstructured":"Figueira MI, Cardoso HJ, Correia S, Maia CJ, Socorro S. The stem cell factor (SCF)\/c-KIT system in carcinogenesis of reproductive tissues: What does the hormonal regulation tell us? Cancer Lett. 2017;405:10\u201321.","journal-title":"Cancer Lett"},{"issue":"9","key":"680_CR82","doi-asserted-by":"publisher","first-page":"3819","DOI":"10.1210\/endo.138.9.5368","volume":"138","author":"JA Parrott","year":"1997","unstructured":"Parrott JA, Skinner MK. Direct actions of kit-ligand on theca cell growth and differentiation during follicle development. Endocrinology. 1997;138(9):3819\u201327.","journal-title":"Endocrinology"},{"issue":"2","key":"680_CR83","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1093\/molehr\/gal010","volume":"12","author":"KJ Hutt","year":"2006","unstructured":"Hutt KJ, McLaughlin EA, Holland MK. Kit ligand and c-Kit have diverse roles during mammalian oogenesis and folliculogenesis. Mol Hum Reprod. 2006;12(2):61\u20139.","journal-title":"Mol Hum Reprod"},{"issue":"1\u20132","key":"680_CR84","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1016\/S0303-7207(01)00746-8","volume":"188","author":"EE Nilsson","year":"2002","unstructured":"Nilsson EE, Kezele P, Skinner MK. Leukemia inhibitory factor (LIF) promotes the primordial to primary follicle transition in rat ovaries. Mol Cell Endocrinol. 2002;188(1\u20132):65\u201373.","journal-title":"Mol Cell Endocrinol"},{"key":"680_CR85","unstructured":"Chansaenroj A. Follicular responsiveness to equine chorionic gonadotropin in cat species: study in in vitro culture. Vet Sci. 2018."},{"issue":"11","key":"680_CR86","doi-asserted-by":"publisher","first-page":"971","DOI":"10.1080\/09513590.2022.2125951","volume":"38","author":"L-Y Cao","year":"2022","unstructured":"Cao L-Y, Zhang Z-Q, Liu P-P, Xu D-F, Tang L, Fan L, et al. Aberrant BMP15\/HIF-1$\/SCF signaling pathway in human granulosa cells is involved in the PCOS related abnormal follicular development. Gynaecol Endocrinol. 2022;38(11):971\u20137.","journal-title":"Gynaecol Endocrinol"},{"key":"680_CR87","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.theriogenology.2021.09.008","volume":"176","author":"C Guo","year":"2021","unstructured":"Guo C, Dong J, Ma Y, Zhou S, Zeng W, Liu G, et al. LIF and bFGF enhanced chicken primordial follicle activation by Wnt\/#-catenin pathway. Theriogenology. 2021;176:1\u201311.","journal-title":"Theriogenology"},{"issue":"5","key":"680_CR88","doi-asserted-by":"publisher","first-page":"967","DOI":"10.1095\/biolreprod.105.043117","volume":"73","author":"P Kezele","year":"2005","unstructured":"Kezele P, Nilsson EE, Skinner MK. Keratinocyte growth factor acts as a mesenchymal factor that promotes ovarian primordial to primary follicle transition. Biol Reprod. 2005;73(5):967\u201373.","journal-title":"Biol Reprod"},{"issue":"7","key":"680_CR89","doi-asserted-by":"publisher","first-page":"905","DOI":"10.1071\/RD11318","volume":"24","author":"RN de Chaves","year":"2012","unstructured":"Chaves RN, de Matos MHT, Buratini J, de Figueiredo JR. The fibroblast growth factor family: involvement in the regulation of folliculogenesis. Reprod Fertil Dev. 2012;24(7):905\u201315.","journal-title":"Reprod Fertil Dev"},{"issue":"1","key":"680_CR90","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1530\/JOE-12-0293","volume":"215","author":"N Schreiber","year":"2012","unstructured":"Schreiber N, Totty M, Spicer L. Expression and effect of fibroblast growth factor 9 in bovine theca cells. J Endocrinol. 2012;215(1):167\u201375.","journal-title":"J Endocrinol"},{"issue":"8","key":"680_CR91","doi-asserted-by":"publisher","first-page":"3711","DOI":"10.1210\/en.2006-1668","volume":"148","author":"AE Drummond","year":"2007","unstructured":"Drummond AE, Tellbach M, Dyson M, Findlay JK. Fibroblast growth factor-9, a local regulator of ovarian function. Endocrinology. 2007;148(8):3711\u201321.","journal-title":"Endocrinology"},{"issue":"9","key":"680_CR92","doi-asserted-by":"publisher","first-page":"4491","DOI":"10.1210\/en.2012-1003","volume":"153","author":"NB Schreiber","year":"2012","unstructured":"Schreiber NB, Spicer LJ. Effects of fibroblast growth factor 9 (FGF9) on steroidogenesis and gene expression and control of FGF9 mRNA in bovine granulosa cells. Endocrinology. 2012;153(9):4491\u2013501.","journal-title":"Endocrinology"},{"issue":"3","key":"680_CR93","doi-asserted-by":"publisher","first-page":"1018","DOI":"10.1095\/biolreprod.101.002527","volume":"67","author":"EE Nilsson","year":"2002","unstructured":"Nilsson EE, Skinner MK. Growth and differentiation factor-9 stimulates progression of early primary but not primordial rat ovarian follicle development. Biol Reprod. 2002;67(3):1018\u201324.","journal-title":"Biol Reprod"},{"issue":"6","key":"680_CR94","doi-asserted-by":"publisher","first-page":"1018","DOI":"10.1210\/mend.13.6.0309","volume":"13","author":"JA Elvin","year":"1999","unstructured":"Elvin JA, Yan C, Wang P, Nishimori K, Matzuk MM. Molecular characterization of the follicle defects in the growth differentiation factor 9-deficient ovary. Mol Endocrinol. 1999;13(6):1018\u201334.","journal-title":"Mol Endocrinol"},{"issue":"2","key":"680_CR95","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1530\/rep.1.01074","volume":"132","author":"PG Knight","year":"2006","unstructured":"Knight PG, Glister C. TGF-# superfamily members and ovarian follicle development. Reproduction. 2006;132(2):191\u2013206.","journal-title":"Reproduction"},{"key":"680_CR96","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13048-018-0392-1","volume":"11","author":"RR de Mahamed","year":"2018","unstructured":"Mahamed RR, Maganhin CC, Sasso GRS, de Jesus Sim\u00f5es M, Baracat MCP, Baracat EC, et al. Metformin improves ovarian follicle dynamics by reducing theca cell proliferation and CYP-17 expression in an androgenized rat model. J Ovarian Res. 2018;11:1\u201310.","journal-title":"J Ovarian Res"},{"issue":"1","key":"680_CR97","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1530\/JME-11-0134","volume":"48","author":"J Young","year":"2011","unstructured":"Young J, McNeilly A. Inhibin removes the inhibitory effects of activin on steroid enzyme expression and androgen production by normal ovarian thecal cells. J Mol Endocrinol. 2011;48(1):49.","journal-title":"J Mol Endocrinol"},{"issue":"2","key":"680_CR98","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1677\/joe.0.1480213","volume":"148","author":"C Sawetawan","year":"1996","unstructured":"Sawetawan C, Carr B, McGee E, Bird I, Hong T, Rainey W. Inhibin and activin differentially regulate androgen production and 17$-hydroxylase expression in human ovarian thecal-like tumor cells. J Endocrinol. 1996;148(2):213\u201321.","journal-title":"J Endocrinol"},{"issue":"1","key":"680_CR99","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1152\/physrev.00002.2018","volume":"99","author":"E Bloise","year":"2019","unstructured":"Bloise E, Ciarmela P, Dela Cruz C, Luisi S, Petraglia F, Reis FM. Activin A in mammalian physiology. Physiol Rev. 2019;99(1):739\u201380.","journal-title":"Physiol Rev"},{"issue":"5","key":"680_CR100","doi-asserted-by":"publisher","first-page":"1151","DOI":"10.1093\/humrep\/den070","volume":"23","author":"EE Telfer","year":"2008","unstructured":"Telfer EE, McLaughlin M, Ding C, Thong KJ. A two-step serum-free culture system supports development of human oocytes from primordial follicles in the presence of activin. Hum Reprod. 2008;23(5):1151\u20138.","journal-title":"Hum Reprod"},{"issue":"3","key":"680_CR101","doi-asserted-by":"publisher","first-page":"967","DOI":"10.1095\/biolreprod65.3.967","volume":"65","author":"J van der Zhao","year":"2001","unstructured":"Zhao J, Taverne MA, van der Weijden GC, Bevers MM, van den Hurk R. Effect of activin A on in vitro development of rat preantral follicles and localization of activin A and activin receptor II. Biol Reprod. 2001;65(3):967\u201377.","journal-title":"Biol Reprod"},{"issue":"1\u20132","key":"680_CR102","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/0303-7207(93)90207-Z","volume":"97","author":"L Shukovski","year":"1993","unstructured":"Shukovski L, Dyson M, Findlay JK. The effects of follistatin, activin and inhibin on steroidogenesis by bovine thecal cells. Mol Cell Endocrinol. 1993;97(1\u20132):19\u201327.","journal-title":"Mol Cell Endocrinol"},{"issue":"6","key":"680_CR103","doi-asserted-by":"publisher","first-page":"1206","DOI":"10.1210\/jcem-72-6-1206","volume":"72","author":"S Hillier","year":"1991","unstructured":"Hillier S, Yong E, Illingworth P, Baird D, Schwall R, Mason A. Effect of recombinant activin on androgen synthesis in cultured human thecal cells. J Clin Endocrinol Metab. 1991;72(6):1206\u201311.","journal-title":"J Clin Endocrinol Metab"},{"issue":"3","key":"680_CR104","doi-asserted-by":"publisher","first-page":"704","DOI":"10.1095\/biolreprod65.3.704","volume":"65","author":"AJ Duleba","year":"2001","unstructured":"Duleba AJ, Pehlivan T, Carbone R, Spaczynski RZ. Activin stimulates proliferation of rat ovarian thecal-interstitial cells. Biol Reprod. 2001;65(3):704\u20139.","journal-title":"Biol Reprod"},{"issue":"10","key":"680_CR105","doi-asserted-by":"publisher","first-page":"9135","DOI":"10.1074\/jbc.M409486200","volume":"280","author":"Y Park","year":"2005","unstructured":"Park Y, Maizels ET, Feiger ZJ, Alam H, Peters CA, Woodruff TK, et al. Induction of cyclin D2 in rat granulosa cells requires FSH-dependent relief from FOXO1 repression coupled with positive signals from Smad. J Biol Chem. 2005;280(10):9135\u201348.","journal-title":"J Biol Chem"},{"issue":"1\u20132","key":"680_CR106","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1016\/j.mce.2003.08.008","volume":"210","author":"T Ogawa","year":"2003","unstructured":"Ogawa T, Yogo K, Ishida N, Takeya T. Synergistic effects of activin and FSH on hyperphosphorylation of Rb and G1\/S transition in rat primary granulosa cells. Mol Cell Endocrinol. 2003;210(1\u20132):31\u20138.","journal-title":"Mol Cell Endocrinol"},{"issue":"7","key":"680_CR107","doi-asserted-by":"publisher","first-page":"a021881","DOI":"10.1101\/cshperspect.a021881","volume":"8","author":"M Namwanje","year":"2016","unstructured":"Namwanje M, Brown CW. Activins and inhibins: roles in development, physiology, and disease. Cold Spring Harb Perspect Biol. 2016;8(7):a021881.","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"680_CR108","doi-asserted-by":"publisher","unstructured":"van Den Driesche S. Activin A reduces luteinisation of human luteinised granulosa cells and has opposing effects to human chorionic gonadotropin in vitro. J Endocrinol. 2008.https:\/\/doi.org\/10.1677\/JOE-08-0302","DOI":"10.1677\/JOE-08-0302"},{"issue":"1","key":"680_CR109","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1210\/er.2003-0007","volume":"25","author":"S Shimasaki","year":"2004","unstructured":"Shimasaki S, Moore RK, Otsuka F, Erickson GF. The bone morphogenetic protein system in mammalian reproduction. Endocr Rev. 2004;25(1):72\u2013101.","journal-title":"Endocr Rev"},{"issue":"6","key":"680_CR110","doi-asserted-by":"publisher","first-page":"980","DOI":"10.3390\/ani13060980","volume":"13","author":"J Jitjumnong","year":"2023","unstructured":"Jitjumnong J, Tang P-C. Bone morphogenetic protein 15 (BMP-15) improves in vitro mouse folliculogenesis. Animals. 2023;13(6):980.","journal-title":"Animals"},{"key":"680_CR111","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1002\/mrd.20163","volume":"69","author":"TK Lwysy","year":"2004","unstructured":"Lwysy TK, Cha KY, Lee SH, Shimasaki S, Lee S, Lee KA. Effects of bone morphogenetic protein-7 (BMP-7) on primordial follicular growth in the mouse ovary. Mol Reprod Dev. 2004;69:159\u201363.","journal-title":"Mol Reprod Dev"},{"issue":"9","key":"680_CR112","doi-asserted-by":"publisher","first-page":"3331","DOI":"10.1210\/jcem.85.9.6835","volume":"85","author":"CA Dooley","year":"2000","unstructured":"Dooley CA, Attia GR, Rainey WE, Moore DR, Carr BR. Bone morphogenetic protein inhibits ovarian androgen production. J Clin Endocrinol Metab. 2000;85(9):3331\u20137.","journal-title":"J Clin Endocrinol Metab"},{"issue":"13","key":"680_CR113","doi-asserted-by":"publisher","first-page":"3775","DOI":"10.3390\/jcm13133775","volume":"13","author":"S Fountas","year":"2024","unstructured":"Fountas S, Petinaki E, Bolaris S, Kargakou M, Dafopoulos S, Zikopoulos A, et al. The roles of GDF-9, BMP-15, BMP-4 and EMMPRIN in folliculogenesis and in vitro fertilization. J Clin Med. 2024;13(13):3775.","journal-title":"J Clin Med"},{"issue":"4","key":"680_CR114","doi-asserted-by":"publisher","first-page":"1608","DOI":"10.1210\/en.2005-0604","volume":"147","author":"B Campbell","year":"2006","unstructured":"Campbell B, Souza C, Skinner A, Webb R, Baird D. Enhanced response of granulosa and theca cells from sheep carriers of the FecB mutation in vitro to gonadotropins and bone morphogenic protein-2,-4, and-6. Endocrinology. 2006;147(4):1608\u201320.","journal-title":"Endocrinology"},{"issue":"4","key":"680_CR115","doi-asserted-by":"publisher","first-page":"1883","DOI":"10.1210\/en.2004-1303","volume":"146","author":"C Glister","year":"2005","unstructured":"Glister C, Richards SL, Knight PG. Bone morphogenetic proteins (BMP)-4,-6, and-7 potently suppress basal and luteinizing hormone-induced androgen production by bovine theca interna cells in primary culture: Could ovarian hyperandrogenic dysfunction be caused by a defect in thecal BMP. signaling? Endocrinol. 2005;146(4):1883\u201392.","journal-title":"signaling? Endocrinol"},{"issue":"1","key":"680_CR116","doi-asserted-by":"publisher","first-page":"S25","DOI":"10.1186\/1297-9686-37-S1-S25","volume":"37","author":"KP McNatty","year":"2005","unstructured":"McNatty KP, Galloway SM, Wilson T, Smith P, Hudson NL, O\u2019Connell A, et al. Physiological effects of major genes affecting ovulation rate in sheep. Genet Select Evol. 2005;37(1):S25\u201338.","journal-title":"Genet Select Evol"},{"issue":"1","key":"680_CR117","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1186\/s12958-022-00913-6","volume":"20","author":"F Afkhami","year":"2022","unstructured":"Afkhami F, Shahbazi S, Farzadi L, Danaei S. Novel bone morphogenetic protein 15 (BMP15) gene variants implicated in premature ovarian insufficiency. Reprod Biol Endocrinol. 2022;20(1):42.","journal-title":"Reprod Biol Endocrinol"},{"issue":"6","key":"680_CR118","doi-asserted-by":"publisher","first-page":"869","DOI":"10.1093\/humupd\/dmu036","volume":"20","author":"L Di Persani","year":"2014","unstructured":"Persani L, Rossetti R, Di Pasquale E, Cacciatore C, Fabre S. The fundamental role of bone morphogenetic protein 15 in ovarian function and its involvement in female fertility disorders. Hum Reprod Update. 2014;20(6):869\u201383.","journal-title":"Hum Reprod Update"},{"issue":"10","key":"680_CR119","doi-asserted-by":"publisher","first-page":"1741","DOI":"10.1007\/s10815-018-1268-4","volume":"35","author":"A Sanfins","year":"2018","unstructured":"Sanfins A, Rodrigues P, Albertini DF. GDF-9 and BMP-15 direct the follicle symphony. J Assist Reprod Genet. 2018;35(10):1741\u201350.","journal-title":"J Assist Reprod Genet"},{"issue":"3","key":"680_CR120","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1262\/jrd.2021-040","volume":"67","author":"G Nagamatsu","year":"2021","unstructured":"Nagamatsu G. Regulation of primordial follicle formation, dormancy, and activation in mice. J Reprod Dev. 2021;67(3):189\u201395.","journal-title":"J Reprod Dev"},{"issue":"1","key":"680_CR121","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1186\/s12964-024-01616-8","volume":"22","author":"Y Duan","year":"2024","unstructured":"Duan Y, Cai B, Guo J, Wang C, Mai Q, Xu Y, et al. GDF9 His209GlnfsTer6\/S428T and GDF9 Q321X\/S428T bi-allelic variants caused female subfertility with defective follicle enlargement. Cell Commun Signal. 2024;22(1):235.","journal-title":"Cell Commun Signal"},{"issue":"2","key":"680_CR122","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1095\/biolreprod.107.063446","volume":"78","author":"LJ Spicer","year":"2008","unstructured":"Spicer LJ, Aad PY, Allen DT, Mazerbourg S, Payne AH, Hsueh AJ. Growth differentiation factor 9 (GDF9) stimulates proliferation and inhibits steroidogenesis by bovine theca cells: influence of follicle size on responses to GDF9. Biol Reprod. 2008;78(2):243\u201353.","journal-title":"Biol Reprod"},{"issue":"6","key":"680_CR123","doi-asserted-by":"publisher","first-page":"921","DOI":"10.1530\/REP-09-0193","volume":"138","author":"L Nicol","year":"2009","unstructured":"Nicol L, Bishop SC, Pong-Wong R, Bendixen C, Holm L-E, Rhind SM, et al. Homozygosity for a single base-pair mutation in the oocyte-specific GDF9 gene results in sterility in Thoka sheep. Reproduction. 2009;138(6):921.","journal-title":"Reproduction"},{"key":"680_CR124","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1016\/bs.vh.2017.12.003","volume":"107","author":"M Belli","year":"2018","unstructured":"Belli M, Shimasaki S. Molecular aspects and clinical relevance of GDF9 and BMP15 in ovarian function. Vitam Horm. 2018;107:317\u201348.","journal-title":"Vitam Horm"},{"issue":"3","key":"680_CR125","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1095\/biolreprod.103.023234","volume":"70","author":"J Wang","year":"2004","unstructured":"Wang J, Roy SK. Growth differentiation factor-9 and stem cell factor promote primordial follicle formation in the hamster: modulation by follicle-stimulating hormone. Biol Reprod. 2004;70(3):577\u201385.","journal-title":"Biol Reprod"},{"issue":"10","key":"680_CR126","doi-asserted-by":"publisher","first-page":"3220","DOI":"10.3390\/ijms19103220","volume":"19","author":"A Schliermann","year":"2018","unstructured":"Schliermann A, Nickel J. Unraveling the connection between fibroblast growth factor and bone morphogenetic protein signaling. Int J Mol Sci. 2018;19(10):3220.","journal-title":"Int J Mol Sci"},{"key":"680_CR127","doi-asserted-by":"publisher","first-page":"1387","DOI":"10.1007\/s10815-021-02161-w","volume":"38","author":"P Asiabi","year":"2021","unstructured":"Asiabi P, David C, Camboni A, Marbaix E, Dolmans M-M, Amorim CA. New insights into the GDF9-Hedgehog-GLI signaling pathway in human ovaries: from fetus to postmenopause. J Assist Reprod Genet. 2021;38:1387\u2013403.","journal-title":"J Assist Reprod Genet"},{"issue":"5","key":"680_CR128","doi-asserted-by":"publisher","first-page":"711","DOI":"10.1210\/me.2008-0391","volume":"23","author":"Y Ren","year":"2009","unstructured":"Ren Y, Cowan RG, Harman RM, Quirk SM. Dominant activation of the hedgehog signaling pathway in the ovary alters theca development and prevents ovulation. Mol Endocrinol. 2009;23(5):711\u201323.","journal-title":"Mol Endocrinol"},{"issue":"1","key":"680_CR129","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1146\/annurev-physiol-061214-111754","volume":"77","author":"I Finco","year":"2015","unstructured":"Finco I, LaPensee CR, Krill KT, Hammer GD. Hedgehog signaling and steroidogenesis. Annu Rev Physiol. 2015;77(1):105\u201329.","journal-title":"Annu Rev Physiol"},{"issue":"28","key":"680_CR130","doi-asserted-by":"publisher","first-page":"11740","DOI":"10.1074\/jbc.M117.781369","volume":"292","author":"Y Liu","year":"2017","unstructured":"Liu Y, Du S-Y, Ding M, Dou X, Zhang F-F, Wu Z-Y, et al. The BMP4-Smad signaling pathway regulates hyperandrogenism development in a female mouse model. J Biol Chem. 2017;292(28):11740\u201350.","journal-title":"J Biol Chem"},{"issue":"1","key":"680_CR131","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1093\/biolre\/ioz167","volume":"102","author":"P Asiabi","year":"2020","unstructured":"Asiabi P, Leonel E, Marbaix E, Dolmans M-M, Amorim C. Immunodetection and quantification of enzymatic markers in theca cells: the early process of ovarian steroidogenesis. Biol Reprod. 2020;102(1):145\u201355.","journal-title":"Biol Reprod"},{"issue":"1","key":"680_CR132","first-page":"818010","volume":"2014","author":"CG Gerv\u00e1sio","year":"2014","unstructured":"Gerv\u00e1sio CG, Bernuci MP, Silva-de-S\u00e1 MF, Rosa-e-Silva ACJS. The role of androgen hormones in early follicular development. Int Sch Res Notices. 2014;2014(1):818010.","journal-title":"Int Sch Res Notices"},{"issue":"3\u20134","key":"680_CR133","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1016\/S0378-4320(03)00088-5","volume":"78","author":"J Fortune","year":"2003","unstructured":"Fortune J. The early stages of follicular development: activation of primordial follicles and growth of preantral follicles. Anim Reprod Sci. 2003;78(3\u20134):135\u201363.","journal-title":"Anim Reprod Sci"},{"issue":"11","key":"680_CR134","doi-asserted-by":"publisher","first-page":"978","DOI":"10.1177\/1933719110379055","volume":"17","author":"SS Patel","year":"2010","unstructured":"Patel SS, Beshay VE, Escobar JC, Carr BR. 17$-Hydroxylase (CYP17) expression and subsequent androstenedione production in the human ovary. Reprod Sci. 2010;17(11):978\u201386.","journal-title":"Reprod Sci"},{"key":"680_CR135","doi-asserted-by":"publisher","first-page":"404005","DOI":"10.3389\/fendo.2018.00452","volume":"9","author":"S Franks","year":"2018","unstructured":"Franks S, Hardy K. Androgen action in the ovary. Front Endocrinol. 2018;9:404005.","journal-title":"Front Endocrinol"},{"key":"680_CR136","volume-title":"Man and woman: a study of secondary and tertiary sexual characters","author":"H Ellis","year":"2013","unstructured":"Ellis H. Man and woman: a study of secondary and tertiary sexual characters. Amsterdam: Elsevier; 2013."},{"issue":"1","key":"680_CR137","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1152\/physrev.2000.80.1.1","volume":"80","author":"GD Niswender","year":"2000","unstructured":"Niswender GD, Juengel JL, Silva PJ, Rollyson MK, McIntush EW. Mechanisms controlling the function and life span of the corpus luteum. Physiol Rev. 2000;80(1):1\u201329.","journal-title":"Physiol Rev"},{"key":"680_CR138","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12958-018-0445-8","volume":"17","author":"B Pan","year":"2019","unstructured":"Pan B, Li J. The art of oocyte meiotic arrest regulation. Reprod Biol Endocrinol. 2019;17:1\u201312.","journal-title":"Reprod Biol Endocrinol"},{"issue":"5","key":"680_CR139","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1016\/S1472-6483(10)60392-6","volume":"15","author":"K Tajima","year":"2007","unstructured":"Tajima K, Orisaka M, Mori T, Kotsuji F. Ovarian theca cells in follicular function. Reprod Biomed Online. 2007;15(5):591\u2013609.","journal-title":"Reprod Biomed Online"},{"issue":"1","key":"680_CR140","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1111\/rda.12790","volume":"52","author":"E Sinderewicz","year":"2017","unstructured":"Sinderewicz E, Grycmacher K, Boruszewska D, Kowalczyk-Zieba I, Woclawek-Potocka I. Lysophosphatidic acid expression in theca cells depends on the type of bovine ovarian follicle. Reprod Domest Anim. 2017;52(1):28\u201334.","journal-title":"Reprod Domest Anim"},{"issue":"9","key":"680_CR141","doi-asserted-by":"publisher","first-page":"3376","DOI":"10.1210\/jc.2018-00549","volume":"103","author":"KY Cheon","year":"2018","unstructured":"Cheon KY, Chung YJ, Cho HH, Kim MR, Cha JH, Kang CS, et al. Expression of mullerian-inhibiting substance\/anti-mullerian hormone type II receptor in the human theca cells. J Clin Endocrinol Metab. 2018;103(9):3376\u201385.","journal-title":"J Clin Endocrinol Metab"},{"key":"680_CR142","doi-asserted-by":"publisher","first-page":"106216","DOI":"10.1016\/j.jsbmb.2022.106216","volume":"226","author":"M Chen","year":"2023","unstructured":"Chen M, Guo X, Zhong Y, Liu Y, Cai B, Wu R, et al. AMH inhibits androgen production in human theca cells. J Steroid Biochem Mol Biol. 2023;226:106216.","journal-title":"J Steroid Biochem Mol Biol"},{"issue":"7","key":"680_CR143","doi-asserted-by":"publisher","first-page":"e0197894","DOI":"10.1371\/journal.pone.0197894","volume":"13","author":"FR Almeida","year":"2018","unstructured":"Almeida FR, Costermans NG, Soede NM, Bunschoten A, Keijer J, Kemp B, et al. Presence of anti-M\u00fcllerian hormone (AMH) during follicular development in the porcine ovary. PLoS ONE. 2018;13(7):e0197894.","journal-title":"PLoS ONE"},{"key":"680_CR144","first-page":"925691","volume":"2015","author":"X Liu","year":"2015","unstructured":"Liu X, Qiao P, Jiang A, Jiang J, Han H, Wang L, et al. Paracrine regulation of steroidogenesis in theca cells by granulosa cells derived from mouse preantral follicles. Biomed Res Int. 2015;2015:925691.","journal-title":"Biomed Res Int"},{"issue":"1","key":"680_CR145","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1186\/s13756-020-00746-1","volume":"9","author":"X Lai","year":"2020","unstructured":"Lai X, Wang X, Yang Q, Xu X, Tang Y, Liu C, et al. Will healthcare workers improve infection prevention and control behaviors as COVID-19 risk emerges and increases, in China? Antimicrob Resist Infect Control. 2020;9(1):83.","journal-title":"Antimicrob Resist Infect Control"},{"key":"680_CR146","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.theriogenology.2019.09.037","volume":"142","author":"Y Yang","year":"2020","unstructured":"Yang Y, Kanno C, Sakaguchi K, Katagiri S, Yanagawa Y, Nagano M. Theca cells can support bovine oocyte growth in vitro without the addition of steroid hormones. Theriogenology. 2020;142:41\u20137.","journal-title":"Theriogenology"},{"issue":"1","key":"680_CR147","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1677\/joe.1.06689","volume":"192","author":"CR Harlow","year":"2007","unstructured":"Harlow CR, Bradshaw AC, Rae MT, Shearer KD, Hillier SG. Oestrogen formation and connective tissue growth factor expression in rat granulosa cells. J Endocrinol. 2007;192(1):41\u201352.","journal-title":"J Endocrinol"},{"key":"680_CR148","first-page":"40","volume":"27","author":"R Jing","year":"2018","unstructured":"Jing R, Gu L, Li J, Gong Y. A transcriptomic comparison of theca and granulosa cells in chicken and cattle follicles reveals ESR2 as a potential regulator of CYP19A1 expression in the theca cells of chicken follicles. Comp Biochem Physiol D Genom Proteom. 2018;27:40\u201353.","journal-title":"Comp Biochem Physiol D Genom Proteom"},{"issue":"6","key":"680_CR149","doi-asserted-by":"publisher","first-page":"869","DOI":"10.1530\/REP-09-0283","volume":"138","author":"RS Robinson","year":"2009","unstructured":"Robinson RS, Woad KJ, Hammond AJ, Laird M, Hunter MG, Mann GE. Angiogenesis and vascular function in the ovary. Reproduction. 2009;138(6):869\u201381.","journal-title":"Reproduction"},{"issue":"1","key":"680_CR150","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1186\/s13048-017-0308-5","volume":"10","author":"Q Xie","year":"2017","unstructured":"Xie Q, Cheng Z, Chen X, Lobe CG, Liu J. The role of Notch signalling in ovarian angiogenesis. J Ovarian Res. 2017;10(1):13.","journal-title":"J Ovarian Res"},{"issue":"4","key":"680_CR151","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1111\/j.1439-0531.2004.00505.x","volume":"39","author":"C De Tamanini","year":"2004","unstructured":"Tamanini C, De Ambrogi M. Angiogenesis in developing follicle and corpus luteum. Reprod Domest Anim. 2004;39(4):206\u201316.","journal-title":"Reprod Domest Anim"},{"issue":"3","key":"680_CR152","doi-asserted-by":"publisher","first-page":"879","DOI":"10.1095\/biolreprod65.3.879","volume":"65","author":"DA Redmer","year":"2001","unstructured":"Redmer DA, Doraiswamy V, Bortnem BJ, Fisher K, Jablonka-Shariff A, Grazul-Bilska AT, et al. Evidence for a role of capillary pericytes in vascular growth of the developing ovine corpus luteum. Biol Reprod. 2001;65(3):879\u201389.","journal-title":"Biol Reprod"},{"issue":"2","key":"680_CR153","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1210\/endo-109-2-356","volume":"109","author":"AJ Zelezni","year":"1981","unstructured":"Zelezni AJ, Schuler HM, Reichert LE. Gonadotropin-binding sites in the rhesus monkey ovary: role of the vasculature in the selective distribution of human chorionic gonadotropin to the preovulatory follicle. Endocrinology. 1981;109(2):356\u201362.","journal-title":"Endocrinology"},{"issue":"1","key":"680_CR154","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1095\/biolreprod41.1.175","volume":"41","author":"GS Greenwald","year":"1989","unstructured":"Greenwald GS. Temporal and topographic changes in DNA synthesis after induced follicular atresia. Biol Reprod. 1989;41(1):175\u201381.","journal-title":"Biol Reprod"},{"issue":"6353","key":"680_CR155","doi-asserted-by":"publisher","first-page":"eaal2379","DOI":"10.1126\/science.aal2379","volume":"357","author":"HG Augustin","year":"2017","unstructured":"Augustin HG, Koh GY. Organotypic vasculature: from descriptive heterogeneity to functional pathophysiology. Science. 2017;357(6353):eaal2379.","journal-title":"Science"},{"issue":"1","key":"680_CR156","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1095\/biolreprod48.1.165","volume":"48","author":"M Br\u00e4nnstr\u00f6m","year":"1993","unstructured":"Br\u00e4nnstr\u00f6m M, Wang L, Norman RJ. Effects of cytokines on prostaglandin production and steroidogenesis of incubated preovulatory follicles of the rat. Biol Reprod. 1993;48(1):165\u201371.","journal-title":"Biol Reprod"},{"issue":"3","key":"680_CR157","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1038\/nri2711","volume":"10","author":"M Saraiva","year":"2010","unstructured":"Saraiva M, O\u2019garra A. The regulation of IL-10 production by immune cells. Nat Rev Immunol. 2010;10(3):170\u201381.","journal-title":"Nat Rev Immunol"},{"issue":"7","key":"680_CR158","first-page":"2414","volume":"84","author":"AJ Jakimiuk","year":"1999","unstructured":"Jakimiuk AJ, Weitsman SR, Magoffin DA. 5$-reductase activity in women with polycystic ovary syndrome. J Clin Endocrinol Metab. 1999;84(7):2414\u20138.","journal-title":"J Clin Endocrinol Metab"},{"issue":"1","key":"680_CR159","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1186\/s13048-025-01837-6","volume":"18","author":"C Zhang","year":"2025","unstructured":"Zhang C, Lin Z, Lin Y, Ma H. Single-cell transcriptomics uncovering a critical AKT-LONP1-STAR axis in ovarian hyperandrogenism of PCOS. J Ovarian Res. 2025;18(1):275.","journal-title":"J Ovarian Res"},{"key":"680_CR160","doi-asserted-by":"crossref","unstructured":"Yuanyuan C, Ningyu S, Lu L, Wuwen Zh, Kai L, Yanting L, Junlan H, Xiang Ch Y, Ping Y. Transcriptomic analysis of theca cells in a PCOS rat model: insights into follicular development. J Ovarian Res. 2025; 18(1):135.","DOI":"10.1186\/s13048-025-01663-w"},{"issue":"1","key":"680_CR161","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1186\/s12958-024-01195-w","volume":"22","author":"S Li","year":"2024","unstructured":"Li S, Li Y, Sun Yu, Feng G, Yang Z, Yan X, Gao X, Jiang Y, Du Y. Zhao Sh, ZhaoH, Chen Z-J. Deconvolution at the single-cell level reveals ovarian cell-type-specific transcriptomic changes in PCOS. Reprod Biol Endocrinol. 2024;22(1):24.","journal-title":"Reprod Biol Endocrinol"},{"issue":"2","key":"680_CR162","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1016\/S0889-8529(05)70074-8","volume":"28","author":"S Franks","year":"1999","unstructured":"Franks S, Gilling-Smith C, Watson H, Willis D. Insulin action in the normal and polycystic ovary. Endocrinol Metab Clin North Am. 1999;28(2):361\u201378.","journal-title":"Endocrinol Metab Clin North Am"},{"issue":"1","key":"680_CR163","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1016\/j.repbio.2015.12.006","volume":"16","author":"D Cadagan","year":"2016","unstructured":"Cadagan D, Khan R, Amer S. Thecal cell sensitivity to luteinizing hormone and insulin in polycystic ovarian syndrome. Reprod Biol. 2016;16(1):53\u201360.","journal-title":"Reprod Biol"},{"key":"680_CR164","doi-asserted-by":"publisher","DOI":"10.1007\/s40618-024-0251","author":"NG Lafci","year":"2024","unstructured":"Lafci NG, Yilmaz B, Yildiz BO. PCOS-the many faces of a disorder in women and men. J Endocrinol Invest. 2024. . https:\/\/doi.org\/10.1007\/s40618-024-0251","journal-title":"J Endocrinol Invest"},{"issue":"6","key":"680_CR165","doi-asserted-by":"publisher","first-page":"981","DOI":"10.1210\/er.2011-1034","volume":"33","author":"E Diamanti-Kandarakis","year":"2012","unstructured":"Diamanti-Kandarakis E, Dunaif A. Insulin resistance and the polycystic ovary syndrome revisited: an update on mechanisms and implications. Endocr Rev. 2012;33(6):981\u20131030.","journal-title":"Endocr Rev"},{"issue":"5","key":"680_CR166","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1210\/er.2015-1104","volume":"37","author":"RL Rosenfield","year":"2016","unstructured":"Rosenfield RL, Ehrmann DA. The pathogenesis of polycystic ovary syndrome (PCOS): the hypothesis of PCOS as functional ovarian hyperandrogenism revisited. Endocr Rev. 2016;37(5):467\u2013520.","journal-title":"Endocr Rev"},{"key":"680_CR167","doi-asserted-by":"publisher","first-page":"1324429","DOI":"10.3389\/fendo.2023.1324429","volume":"14","author":"M Orisaka","year":"2023","unstructured":"Orisaka M, Mizutani T, Miyazaki Y, Shirafuji A, Tamamura C, Fujita M, et al. Chronic low-grade inflammation and ovarian dysfunction in women with polycystic ovarian syndrome, endometriosis, and aging. Front Endocrinol. 2023;14:1324429.","journal-title":"Front Endocrinol"},{"issue":"29","key":"680_CR168","doi-asserted-by":"publisher","first-page":"26380","DOI":"10.1074\/jbc.M300688200","volume":"278","author":"JR Wood","year":"2003","unstructured":"Wood JR, Nelson VL, Ho C, Jansen E, Wang CY, Urbanek M, et al. The molecular phenotype of polycystic ovary syndrome (PCOS) theca cells and new candidate PCOS genes defined by microarray analysis. J Biol Chem. 2003;278(29):26380\u201390.","journal-title":"J Biol Chem"},{"key":"680_CR169","doi-asserted-by":"publisher","first-page":"1273542","DOI":"10.3389\/fendo.2023.1273542","volume":"14","author":"K Wang","year":"2023","unstructured":"Wang K, Li Y, Chen Y. Androgen excess: a hallmark of polycystic ovary syndrome. Front Endocrinol. 2023;14:1273542.","journal-title":"Front Endocrinol"},{"key":"680_CR170","doi-asserted-by":"publisher","DOI":"10.1017\/s1462399408000598","author":"E Diamanti-Kandarakis","year":"2008","unstructured":"Diamanti-Kandarakis E. Polycystic ovarian syndrome: pathophysiology, molecular aspects and clinical implications. Expert Rev Mol Med. 2008. . https:\/\/doi.org\/10.1017\/s1462399408000598","journal-title":"Expert Rev Mol Med"},{"key":"680_CR171","doi-asserted-by":"crossref","unstructured":"Tee MK, Speek M, Legeza B, Modi B, Teves ME, McAllister JM, et al. Alternative splicing of DENND1A, a PCOS candidate gene, generates variant 2. Mol Cell Endocrinol. 2016;434:25\u201335.","DOI":"10.1016\/j.mce.2016.06.011"},{"key":"680_CR172","doi-asserted-by":"publisher","unstructured":"Kobayashi M, Yoshino O, Nakashima A, Ito M, Nishio K, Ono Y et al. Inhibition of autophagy in theca cells induces CYP17A1 and PAI-1 expression via ROS\/p38 and JNK signalling during the development of polycystic ovary syndrome. Mol Cell Endocrinol. 2020. https:\/\/doi.org\/10.1016\/j.mce.2020.110792.","DOI":"10.1016\/j.mce.2020.110792."},{"key":"680_CR173","volume-title":"Seminars in reproductive medicine","author":"R Tal","year":"2015","unstructured":"Tal R, Seifer DB, Arici A. The emerging role of angiogenic factor dysregulation in the pathogenesis of polycystic ovarian syndrome. Seminars in reproductive medicine. Stuttgart: Thieme Medical; 2015."},{"issue":"5","key":"680_CR174","doi-asserted-by":"publisher","first-page":"R199","DOI":"10.1530\/REP-17-0597","volume":"155","author":"M Di Pietro","year":"2018","unstructured":"Di Pietro M, Pascuali N, Parborell F, Abramovich D. Ovarian angiogenesis in polycystic ovary syndrome. Reproduction. 2018;155(5):R199\u2013209.","journal-title":"Reproduction"},{"key":"680_CR175","doi-asserted-by":"publisher","first-page":"857","DOI":"10.1007\/s10815-014-0236-x","volume":"31","author":"KR Reddy","year":"2014","unstructured":"Reddy KR, Deepika M, Supriya K, Latha KP, Rao SL, Rani VU, et al. CYP11A1 microsatellite (tttta) n polymorphism in PCOS women from South India. J Assist Reprod Genet. 2014;31:857\u201363.","journal-title":"J Assist Reprod Genet"},{"key":"680_CR176","doi-asserted-by":"crossref","unstructured":"Rao A, Rao AUJ, Wahab Z, Qudsia HK, Javed R. Association of Cyp11a gene with polycystic ovarian syndrome patients in Lahore, Pakistan: Cyp11a gene with polycystic ovarian syndrome patients. Pak J Health Sci. 2025;325\u201330.","DOI":"10.54393\/pjhs.v6i1.2761"},{"issue":"3","key":"680_CR177","first-page":"245","volume":"22","author":"KM Sulaiman","year":"2024","unstructured":"Sulaiman KM. Study of cytochrome P450 C 17 alpha (CYP17A1) gene polymorphism in patients with poly cystic ovarian syndrome. Gomal J Med Sci. 2024;22(3):245\u201351.","journal-title":"Gomal J Med Sci"},{"issue":"12","key":"680_CR178","doi-asserted-by":"publisher","first-page":"R305","DOI":"10.1530\/EC-21-0327","volume":"10","author":"X Liu","year":"2021","unstructured":"Liu X, Xu M, Qian M, Yang L. CYP17 T\/C (rs74357) gene polymorphism contributes to polycystic ovary syndrome susceptibility: evidence from a meta-analysis. Endocr Connect. 2021;10(12):R305\u201316.","journal-title":"Endocr Connect"},{"issue":"4","key":"680_CR179","first-page":"e2023167","volume":"94","author":"ML El Rezgoun","year":"2023","unstructured":"Rezgoun ML, El Khour D, Bendaoud H, Chellat D. CYP17A1 (rs74357) polymorphism and polycystic ovary syndrome risk: a systemic review and meta-analysis. Acta Bio Med Atenei Parmensis. 2023;94(4):e2023167.","journal-title":"Acta Bio Med Atenei Parmensis"},{"issue":"1","key":"680_CR180","doi-asserted-by":"publisher","first-page":"12942","DOI":"10.1038\/s41598-021-92265-1","volume":"11","author":"S Ashraf","year":"2021","unstructured":"Ashraf S, Rasool SUA, Nabi M, Ganie MA, Masoodi SR, Amin S. Impact of rs2414096 polymorphism of CYP19 gene on susceptibility of polycystic ovary syndrome and hyperandrogenism in Kashmiri women. Sci Rep. 2021;11(1):12942.","journal-title":"Sci Rep"},{"issue":"2","key":"680_CR181","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1080\/09513590.2020.1813274","volume":"37","author":"P Sharma","year":"2021","unstructured":"Sharma P, Kaur M, Khetarpal P. CYP19 gene rs2414096 variant and differential genetic risk of polycystic ovary syndrome: a systematic review and meta-analysis. Gynaecol Endocrinol. 2021;37(2):126\u201331.","journal-title":"Gynaecol Endocrinol"},{"issue":"2","key":"680_CR182","first-page":"149","volume":"39","author":"MH Daghestani","year":"2020","unstructured":"Daghestani MH. Rs1799817 in INSR associates with susceptibility to polycystic ovary syndrome. J Med Biochem. 2020;39(2):149.","journal-title":"J Med Biochem"},{"issue":"5","key":"680_CR183","doi-asserted-by":"publisher","first-page":"1213","DOI":"10.1016\/j.fertnstert.2007.05.026","volume":"89","author":"E-J Lee","year":"2008","unstructured":"Lee E-J, Oh B, Lee J-Y, Kimm K, Lee S-H, Baek K-H. A novel single nucleotide polymorphism of INSR gene for polycystic ovary syndrome. Fertil Steril. 2008;89(5):1213\u201320.","journal-title":"Fertil Steril"},{"issue":"8","key":"680_CR184","doi-asserted-by":"publisher","first-page":"1463","DOI":"10.3390\/genes13081463","volume":"13","author":"SUA Rasool","year":"2022","unstructured":"Rasool SUA, Nabi M, Ashraf S, Amin S. Insulin receptor substrate 1 Gly972Arg (rs1801278) polymorphism is associated with obesity and insulin resistance in Kashmiri women with polycystic ovary syndrome. Genes. 2022;13(8):1463.","journal-title":"Genes"},{"issue":"7","key":"680_CR185","first-page":"844","volume":"42","author":"X Shi","year":"2016","unstructured":"Shi X, Xie X, Jia Y, Li S. Associations of insulin receptor and insulin receptor substrates genetic polymorphisms with polycystic ovary syndrome: a systematic review and meta!analysis. Obstet Gynecol Res. 2016;42(7):844\u201354.","journal-title":"Obstet Gynecol Res"},{"issue":"11","key":"680_CR186","doi-asserted-by":"publisher","first-page":"5529","DOI":"10.1210\/jc.2003-030322","volume":"88","author":"A Gonzalez","year":"2003","unstructured":"Gonzalez A, Abril E, Roca A, Arag\u00f3n MJ, Figueroa MJ, Velarde P, et al. Specific CAPN10 gene haplotypes influence the clinical profile of polycystic ovary patients. J Clin Endocrinol Metab. 2003;88(11):5529\u201336.","journal-title":"J Clin Endocrinol Metab"},{"key":"680_CR187","doi-asserted-by":"publisher","first-page":"1153960","DOI":"10.3389\/fgene.2023.1153960","volume":"14","author":"Y Li","year":"2023","unstructured":"Li Y, Han T, Wang Y, Gao J, Zhang J, Wu Y, et al. Association of Calpain10 polymorphisms with polycystic ovarian syndrome susceptibility: a systematic review and meta-analysis with trial sequential analysis. Front Genet. 2023;14:1153960.","journal-title":"Front Genet"},{"issue":"8","key":"680_CR188","doi-asserted-by":"publisher","first-page":"3971","DOI":"10.1210\/jcem.87.8.8793","volume":"87","author":"A Gonzalez","year":"2002","unstructured":"Gonzalez A, Abril E, Roca A, Arag\u00f3n MJ, Figueroa MJ, Velarde P, et al. CAPN10 alleles are associated with polycystic ovary syndrome. J Clin Endocrinol Metab. 2002;87(8):3971\u20136.","journal-title":"J Clin Endocrinol Metab"},{"issue":"3","key":"680_CR189","first-page":"182","volume":"14","author":"S Beksac","year":"2013","unstructured":"Beksac S, Sel\u00e7uk I, Boyraz G, G\u00fcner G, Turgal M, Usubutun A. Two patients with marginal symptoms showing hyperthecosis at the edge of malignancy: presentation of two cases. J Turk Ger Gynecol Assoc. 2013;14(3):182.","journal-title":"J Turk Ger Gynecol Assoc"},{"issue":"1","key":"680_CR190","first-page":"2783464","volume":"2023","author":"LS Metzker","year":"2023","unstructured":"Metzker LS, Ferreira LAC, Borges JCN, Guzzo MF, Ferreira RN, Silva LLR, et al. Postmenopausal hyperandrogenism due to ovarian hyperthecosis. Case Rep Obstet Gynecol. 2023;2023(1):2783464.","journal-title":"Case Rep Obstet Gynecol"},{"issue":"5","key":"680_CR191","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1007\/s40477-025-01002-1","volume":"28","author":"DL Brown","year":"2009","unstructured":"Brown DL, Henrichsen TL, Clayton AC, Hudson SB, Coddington IIICC, Vella A. Ovarian stromal hyperthecosis: sonographic features and histologic associations. J Ultrasound. 2009;28(5):587\u201393.","journal-title":"J Ultrasound"},{"issue":"6","key":"680_CR192","doi-asserted-by":"publisher","first-page":"luad148","DOI":"10.1210\/jcemcr\/luad148","volume":"1","author":"S Yousaf","year":"2023","unstructured":"Yousaf S, Nizar R, John L, Simpson A. A case of ovarian hyperthecosis in a postmenopausal woman. JCEM Case Rep. 2023;1(6):luad148.","journal-title":"JCEM Case Rep"},{"issue":"2","key":"680_CR193","doi-asserted-by":"publisher","first-page":"384","DOI":"10.1016\/0002-9378(86)90676-9","volume":"154","author":"M Van Nagamani","year":"1986","unstructured":"Nagamani M, Van Dinh T, Kelver ME. Hyperinsulinemia in hyperthecosis of the ovaries. Am J Obstet Gynecol. 1986;154(2):384\u20139.","journal-title":"Am J Obstet Gynecol"},{"issue":"5","key":"680_CR194","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1530\/EJE-11-0542","volume":"165","author":"AP Van Beek","year":"2011","unstructured":"Van Beek AP, Cantineau AE. Ovarian hyperthecosis. Eur J Endocrinol. 2011;165(5):779\u201388.","journal-title":"Eur J Endocrinol"},{"key":"680_CR195","doi-asserted-by":"publisher","first-page":"120","DOI":"10.1016\/j.humpath.2016.09.021","volume":"59","author":"C Zhang","year":"2017","unstructured":"Zhang C, Sung CJ, Quddus MR, Simon RA, Jazaerly T, Lawrence WD. Association of ovarian hyperthecosis with endometrial polyp, endometrial hyperplasia, and endometrioid adenocarcinoma in postmenopausal women: a clinicopathological study of 238 cases. Hum Pathol. 2017;59:120\u20134.","journal-title":"Hum Pathol"},{"issue":"12","key":"680_CR196","doi-asserted-by":"publisher","first-page":"939","DOI":"10.1097\/00000478-198412000-00007","volume":"8","author":"E Ravinsky","year":"1984","unstructured":"Ravinsky E. Ovarian hyperthecosis associated with pseudosarcomatous changes in the endometrial stroma. Am J Surg Pathol. 1984;8(12):939\u201343.","journal-title":"Am J Surg Pathol"},{"issue":"2","key":"680_CR197","doi-asserted-by":"publisher","first-page":"160","DOI":"10.1111\/j.1365-2265.2011.04040.x","volume":"75","author":"MS Rothman","year":"2011","unstructured":"Rothman MS, Wierman ME. How should postmenopausal androgen excess be evaluated? Clin Endocrinol. 2011;75(2):160\u20134.","journal-title":"Clin Endocrinol"},{"issue":"5 Part 2","key":"680_CR198","first-page":"893","volume":"99","author":"E Krug","year":"2002","unstructured":"Krug E, Berga SL. Postmenopausal hyperthecosis: functional dysregulation of androgenesis in climacteric ovary. Obstet Gynecol. 2002;99(5 Part 2):893\u20137.","journal-title":"Obstet Gynecol"},{"key":"680_CR199","doi-asserted-by":"publisher","first-page":"626924","DOI":"10.3389\/fendo.2021.626924","volume":"12","author":"B Ishizuka","year":"2021","unstructured":"Ishizuka B. Current understanding of the etiology, symptomatology, and treatment options in premature ovarian insufficiency (POI). Front Endocrinol. 2021;12:626924.","journal-title":"Front Endocrinol"},{"key":"680_CR200","doi-asserted-by":"publisher","first-page":"1573","DOI":"10.1007\/s10815-014-0342-9","volume":"31","author":"C Fortu\u00f1o","year":"2014","unstructured":"Fortu\u00f1o C, Labarta E. Genetics of primary ovarian insufficiency: a review. J Assist Reprod Genet. 2014;31:1573\u201385.","journal-title":"J Assist Reprod Genet"},{"issue":"2","key":"680_CR201","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1111\/cge.12921","volume":"91","author":"R Rossetti","year":"2017","unstructured":"Rossetti R, Ferrari I, Bonomi M, Persani L. Genetics of primary ovarian insufficiency. Clin Genet. 2017;91(2):183\u201398.","journal-title":"Clin Genet"},{"issue":"1","key":"680_CR202","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1038\/ng0993-83","volume":"5","author":"C Matthews","year":"1993","unstructured":"Matthews C, Borgato S, Beck-Peccoz P, Adams M, Tone Y, Gambino G, et al. Primary amenorrhoea and infertility due to a mutation in the #\u2013subunit of follicle\u2013stimulating hormone. Nat Genet. 1993;5(1):83\u20136.","journal-title":"Nat Genet"},{"key":"680_CR203","doi-asserted-by":"publisher","first-page":"152","DOI":"10.3389\/fendo.2015.00152","volume":"6","author":"P Narayan","year":"2015","unstructured":"Narayan P. Genetic models for the study of luteinizing hormone receptor function. Front Endocrinol. 2015;6:152.","journal-title":"Front Endocrinol"},{"issue":"9","key":"680_CR204","doi-asserted-by":"publisher","first-page":"1206","DOI":"10.1093\/oxfordjournals.humrep.a137513","volume":"6","author":"SC Chappel","year":"1991","unstructured":"Chappel SC, Howles C. Reevaluation of the roles of luteinizing hormone and follicle-stimulating hormone in the ovulatory process. Hum Reprod. 1991;6(9):1206\u201312.","journal-title":"Hum Reprod"},{"key":"680_CR205","doi-asserted-by":"publisher","first-page":"121479","DOI":"10.1016\/j.lfs.2023.121479","volume":"317","author":"H Vlieghe","year":"2023","unstructured":"Vlieghe H, Leonel EC, Asiabi P, Amorim CA. The characterization and therapeutic applications of ovarian theca cells: an update. Life Sci. 2023;317:121479.","journal-title":"Life Sci"},{"issue":"1","key":"680_CR206","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1210\/er.2010-0013","volume":"32","author":"WL Miller","year":"2011","unstructured":"Miller WL, Auchus RJ. The molecular biology, biochemistry, and physiology of human steroidogenesis and its disorders. Endocr Rev. 2011;32(1):81\u2013151.","journal-title":"Endocr Rev"},{"issue":"2","key":"680_CR207","doi-asserted-by":"publisher","first-page":"302","DOI":"10.3390\/genes13020302","volume":"13","author":"R Heidarzadehpilehrood","year":"2022","unstructured":"Heidarzadehpilehrood R, Pirhoushiaran M, Abdollahzadeh R, Binti Osman M, Sakinah M, Nordin N, et al. A review on CYP11A1, CYP17A1, and CYP19A1 polymorphism studies: candidate susceptibility genes for polycystic ovary syndrome (PCOS) and infertility. Genes. 2022;13(2):302.","journal-title":"Genes"},{"issue":"5","key":"680_CR208","doi-asserted-by":"publisher","first-page":"bqaf071","DOI":"10.1210\/endocr\/bqaf071","volume":"166","author":"S Joseph","year":"2025","unstructured":"Joseph S, Ubba V, Wang Z, Feng M, Dsilva MK, Suero S, et al. Ovarian-specific Cyp17A1 overexpression in female mice: a novel model of endogenous testosterone excess. Endocrinology. 2025;166(5):bqaf071.","journal-title":"Endocrinology"},{"issue":"1","key":"680_CR209","doi-asserted-by":"publisher","first-page":"22841","DOI":"10.1038\/s41598-023-48881-0","volume":"13","author":"S Singh","year":"2023","unstructured":"Singh S, Kaur M, Beri A, Kaur A. Significance of LHCGR polymorphisms in polycystic ovary syndrome: an association study. Sci Rep. 2023;13(1):22841.","journal-title":"Sci Rep"},{"issue":"4","key":"680_CR210","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1093\/humupd\/dmp011","volume":"15","author":"R Ivell","year":"2009","unstructured":"Ivell R, Anand-Ivell R. Biology of insulin-like factor 3 in human reproduction. Hum Reprod Update. 2009;15(4):463\u201376.","journal-title":"Hum Reprod Update"},{"issue":"3","key":"680_CR211","doi-asserted-by":"publisher","first-page":"480","DOI":"10.1095\/biolreprod.109.077552","volume":"81","author":"R Anand-Ivell","year":"2009","unstructured":"Anand-Ivell R, Heng K, Hafen B, Setchell B, Ivell R. Dynamics of INSL3 peptide expression in the rodent testis. Biol Reprod. 2009;81(3):480\u20137.","journal-title":"Biol Reprod"},{"issue":"2","key":"680_CR212","doi-asserted-by":"publisher","first-page":"328","DOI":"10.1016\/j.fertnstert.2020.09.145","volume":"115","author":"RW Rebar","year":"2021","unstructured":"Rebar RW, Keator CS. Expanding our knowledge of premature ovarian insufficiency. Fertil Steril. 2021;115(2):328\u20139.","journal-title":"Fertil Steril"},{"issue":"11","key":"680_CR213","doi-asserted-by":"publisher","first-page":"1483","DOI":"10.1007\/s10815-014-0319-8","volume":"31","author":"L-N Wei","year":"2014","unstructured":"Wei L-N, Huang R, Li L-L, Fang C, Li Y, Liang X-Y. Reduced and delayed expression of GDF9 and BMP15 in ovarian tissues from women with polycystic ovary syndrome. J Assist Reprod Genet. 2014;31(11):1483\u201390.","journal-title":"J Assist Reprod Genet"},{"issue":"8","key":"680_CR214","doi-asserted-by":"publisher","first-page":"2174","DOI":"10.1007\/s43032-020-00409-x","volume":"28","author":"A Monte","year":"2021","unstructured":"Monte A, Bezerra M, Menezes V, Gouveia B, Barberino R, Lins T, et al. Involvement of phosphorylated Akt and FOXO3a in the effects of growth and differentiation Factor-9 (GDF-9) on inhibition of follicular apoptosis and induction of granulosa cell proliferation after in vitro culture of sheep ovarian tissue. Reprod Sci. 2021;28(8):2174\u201385.","journal-title":"Reprod Sci"},{"issue":"6","key":"680_CR215","doi-asserted-by":"publisher","first-page":"1117","DOI":"10.1093\/humrep\/deu076","volume":"29","author":"R Homburg","year":"2014","unstructured":"Homburg R, Crawford G. The role of AMH in anovulation associated with PCOS: a hypothesis. Hum Reprod. 2014;29(6):1117\u201321.","journal-title":"Hum Reprod"},{"issue":"3","key":"680_CR216","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1530\/JOE-18-0678","volume":"240","author":"J Yu","year":"2019","unstructured":"Yu J, Liu Y, Zhang D, Zhai D, Song L, Cai Z, et al. Baicalin inhibits recruitment of GATA1 to the HSD3B2 promoter and reverses hyperandrogenism of PCOS. J Endocrinol. 2019;240(3):497\u2013507.","journal-title":"J Endocrinol"},{"issue":"2","key":"680_CR217","doi-asserted-by":"publisher","first-page":"783","DOI":"10.1210\/jc.2003-030934","volume":"89","author":"G Carbunaru","year":"2004","unstructured":"Carbunaru G, Prasad P, Scoccia B, Shea P, Hopwood N, Ziai F, et al. The hormonal phenotype of nonclassic 3#-hydroxysteroid dehydrogenase (HSD3B) deficiency in hyperandrogenic females is associated with insulin-resistant polycystic ovary syndrome and is not a variant of inherited HSD3B2 deficiency. J Clin Endocrinol Metab. 2004;89(2):783\u201394.","journal-title":"J Clin Endocrinol Metab"},{"issue":"2","key":"680_CR218","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1007\/s10815-018-1335-x","volume":"36","author":"Y Geng","year":"2019","unstructured":"Geng Y, Sui C, Xun Y, Lai Q, Jin L. MiRNA-99a can regulate proliferation and apoptosis of human granulosa cells via targeting IGF-1R in polycystic ovary syndrome. J Assist Reprod Genet. 2019;36(2):211\u201321.","journal-title":"J Assist Reprod Genet"},{"issue":"7","key":"680_CR219","doi-asserted-by":"publisher","first-page":"2309","DOI":"10.1210\/en.2017-00146","volume":"158","author":"SC Baumgarten","year":"2017","unstructured":"Baumgarten SC, Armouti M, Ko C, Stocco C. IGF1R expression in ovarian granulosa cells is essential for steroidogenesis, follicle survival, and fertility in female mice. Endocrinology. 2017;158(7):2309\u201318.","journal-title":"Endocrinology"},{"issue":"1","key":"680_CR220","doi-asserted-by":"publisher","first-page":"8993","DOI":"10.1038\/s41598-020-65890-5","volume":"10","author":"X-y Gao","year":"2020","unstructured":"Gao X-y, Liu Y, Lv Y, Huang T, Lu G, Liu H-b, et al. Role of androgen receptor for reconsidering the true polycystic ovarian morphology in PCOS. Sci Rep. 2020;10(1):8993.","journal-title":"Sci Rep"},{"issue":"2","key":"680_CR221","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1016\/j.ijgo.2012.08.016","volume":"120","author":"LH Lin","year":"2013","unstructured":"Lin LH, Baracat MC, Maciel GA, Soares JJM, Baracat EC. Androgen receptor gene polymorphism and polycystic ovary syndrome. Int J Gynaecol Obstet. 2013;120(2):115\u20138.","journal-title":"Int J Gynaecol Obstet"},{"issue":"11","key":"680_CR222","doi-asserted-by":"publisher","first-page":"1318","DOI":"10.3390\/cells8111318","volume":"8","author":"J-F Chen","year":"2019","unstructured":"Chen J-F, Lin P-W, Tsai Y-R, Yang Y-C, Kang H-Y. Androgens and androgen receptor actions on bone health and disease: from androgen deficiency to androgen therapy. Cells. 2019;8(11):1318.","journal-title":"Cells"},{"issue":"11","key":"680_CR223","doi-asserted-by":"publisher","first-page":"e0165636","DOI":"10.1371\/journal.pone.0165636","volume":"11","author":"WY Almawi","year":"2016","unstructured":"Almawi WY, Gammoh E, Malalla ZH, Al-Madhi SA. Analysis of VEGFA variants and changes in VEGF levels underscores the contribution of VEGF to polycystic ovary syndrome. PLoS ONE. 2016;11(11):e0165636.","journal-title":"PLoS ONE"},{"issue":"9","key":"680_CR224","doi-asserted-by":"publisher","first-page":"748","DOI":"10.1186\/s12864-016-3092-5","volume":"17","author":"A Ben Salem","year":"2016","unstructured":"Ben Salem A, Megdich F, Kacem O, Souayeh M, Hachani Ben Ali F, Hizem S, et al. Vascular endothelial growth factor (VEGFA) gene variation in polycystic ovary syndrome in a Tunisian women population. BMC Genom. 2016;17(9):748.","journal-title":"BMC Genom"},{"issue":"1","key":"680_CR225","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1177\/1933719113485294","volume":"21","author":"N Raja-Khan","year":"2014","unstructured":"Raja-Khan N, Urbanek M, Rodgers RJ, Legro RS. The role of TGF-# in polycystic ovary syndrome. Reprod Sci. 2014;21(1):20\u201331.","journal-title":"Reprod Sci"},{"key":"680_CR226","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1016\/j.cca.2020.07.032","volume":"510","author":"M Bhati","year":"2020","unstructured":"Bhati M, Prabhu YD, Renu K, Vellingiri B, Thiagarajan P, Panda A, et al. Role of TGF-# signalling in PCOS associated focal segmental glomerulosclerosis. Clin Chim Acta. 2020;510:244\u201351.","journal-title":"Clin Chim Acta"},{"issue":"1","key":"680_CR227","doi-asserted-by":"publisher","first-page":"6","DOI":"10.1186\/s13048-017-0375-7","volume":"11","author":"D Wang","year":"2018","unstructured":"Wang D, Wang W, Liang Q, He X, Xia Y, Shen S, et al. DHEA-induced ovarian hyperfibrosis is mediated by TGF-# signaling pathway. J Ovarian Res. 2018;11(1):6.","journal-title":"J Ovarian Res"},{"issue":"1","key":"680_CR228","doi-asserted-by":"publisher","first-page":"82","DOI":"10.3109\/13697137.2011.571321","volume":"15","author":"K Ethun Jr","year":"2012","unstructured":"Ethun K, Wood C, Parker C Jr, Kaplan J, Chen H, Appt S. Effect of ovarian aging on androgen biosynthesis in a cynomolgus macaque model. Climacteric. 2012;15(1):82\u201392.","journal-title":"Climacteric"},{"issue":"2","key":"680_CR229","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1038\/s43587-024-00762-5","volume":"5","author":"C Jin","year":"2025","unstructured":"Jin C, Wang X, Yang J, Kim S, Hudgins AD, Gamliel A, et al. Molecular and genetic insights into human ovarian aging from single-nuclei multi-omics analyses. Nat Aging. 2025;5(2):275\u201390.","journal-title":"Nat Aging"},{"key":"680_CR230","doi-asserted-by":"crossref","unstructured":"Jin C, Wang X, Hudgins AD, Gamliel A, Pei M, Kim S et al. The regulatory landscapes of human ovarian ageing. BioRxiv. 2022:2022.05.18.492547.","DOI":"10.1101\/2022.05.18.492547"},{"issue":"2","key":"680_CR231","doi-asserted-by":"publisher","first-page":"R19","DOI":"10.1530\/REP-21-0022","volume":"162","author":"SU Park","year":"2021","unstructured":"Park SU, Walsh L, Berkowitz KM. Mechanisms of ovarian aging. Reproduction. 2021;162(2):R19\u201333.","journal-title":"Reproduction"},{"issue":"1","key":"680_CR232","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1093\/humupd\/dms043","volume":"19","author":"SM Nelson","year":"2013","unstructured":"Nelson SM, Telfer EE, Anderson RA. The ageing ovary and uterus: new biological insights. Hum Reprod Update. 2013;19(1):67\u201383.","journal-title":"Hum Reprod Update"},{"issue":"18","key":"680_CR233","doi-asserted-by":"publisher","first-page":"9906","DOI":"10.3390\/ijms25189906","volume":"25","author":"V la D\u00edaz-Hern\u00e1ndez","year":"2024","unstructured":"D\u00edaz-Hern\u00e1ndez V, Marmolejo-Valencia A, la Montiel-De Cruz C, Pi\u00f1\u00f3n-Z\u00e1rate G, Monta\u00f1o LM, Mora-Herrera SI, et al. The interstitial gland as a source of pro-or anti-senescent cells during Chinchilla rabbit ovarian aging. Int J Mol Sci. 2024;25(18):9906.","journal-title":"Int J Mol Sci"},{"issue":"3","key":"680_CR234","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1097\/GME.0000000000000755","volume":"24","author":"M Pertynska-Marczewska","year":"2017","unstructured":"Pertynska-Marczewska M, Diamanti-Kandarakis E. Aging ovary and the role for advanced glycation end products. Menopause. 2017;24(3):345\u201351.","journal-title":"Menopause"},{"issue":"5","key":"680_CR235","doi-asserted-by":"publisher","first-page":"1546","DOI":"10.1016\/j.fertnstert.2007.06.032","volume":"89","author":"J Yeh","year":"2008","unstructured":"Yeh J, Kim BS, Peresie J. Ovarian vascular endothelial growth factor and vascular endothelial growth factor receptor patterns in reproductive aging. Fertil Steril. 2008;89(5):1546\u201356.","journal-title":"Fertil Steril"},{"issue":"1","key":"680_CR236","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1016\/j.mce.2008.11.004","volume":"299","author":"A Perheentupa","year":"2009","unstructured":"Perheentupa A, Huhtaniemi I. Aging of the human ovary and testis. Mol Cell Endocrinol. 2009;299(1):2\u201313.","journal-title":"Mol Cell Endocrinol"},{"key":"680_CR237","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/1423-0127-20-93","volume":"20","author":"JH Ford","year":"2013","unstructured":"Ford JH. Reduced quality and accelerated follicle loss with female reproductive aging-does decline in theca dehydroepiandrosterone (DHEA) underlie the problem? J Biomed Sci. 2013;20:1\u20139.","journal-title":"J Biomed Sci"},{"issue":"2","key":"680_CR238","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1093\/humupd\/dmm048","volume":"14","author":"C Tatone","year":"2008","unstructured":"Tatone C, Amicarelli F, Carbone MC, Monteleone P, Caserta D, Marci R, et al. Cellular and molecular aspects of ovarian follicle ageing. Hum Reprod Update. 2008;14(2):131\u201342.","journal-title":"Hum Reprod Update"},{"issue":"12","key":"680_CR239","doi-asserted-by":"publisher","first-page":"4835","DOI":"10.1210\/en.2012-2140","volume":"154","author":"PT Jimenez","year":"2013","unstructured":"Jimenez PT, Frolova AI, Chi MM, Grindler NM, Willcockson AR, Reynolds KA, et al. DHEA-mediated inhibition of the pentose phosphate pathway alters oocyte lipid metabolism in mice. Endocrinology. 2013;154(12):4835\u201344.","journal-title":"Endocrinology"},{"key":"680_CR240","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1016\/bs.vh.2018.02.001","volume":"108","author":"BN Chimote","year":"2018","unstructured":"Chimote BN, Chimote NM. Dehydroepiandrosterone (DHEA) and its sulfate (DHEA-S) in mammalian reproduction: known roles and novel paradigms. Vitam Horm. 2018;108:223\u201350.","journal-title":"Vitam Horm"},{"issue":"1","key":"680_CR241","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/S0026-895X(25)09162-X","volume":"50","author":"JM Peters","year":"1996","unstructured":"Peters JM, Zhou Y-C, Ram PA, Lee S, Gonzalez FJ, Waxman DJ. Peroxisome proliferator-activated receptor alpha required for gene induction by dehydroepiandrosterone-3 beta-sulfate. Mol Pharmacol. 1996;50(1):67\u201374.","journal-title":"Mol Pharmacol"},{"issue":"1","key":"680_CR242","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1210\/en.2002-220039","volume":"144","author":"K Kajita","year":"2003","unstructured":"Kajita K, Ishizuka T, Mune T, Miura A, Ishizawa M, Kanoh Y, et al. Dehydroepiandrosterone down-regulates the expression of peroxisome proliferatoractivated receptor * in adipocytes. Endocrinology. 2003;144(1):253\u20139.","journal-title":"Endocrinology"}],"container-title":["Biological Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s40659-026-00680-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s40659-026-00680-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s40659-026-00680-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T23:03:30Z","timestamp":1790982210000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s40659-026-00680-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,8]]},"references-count":242,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["680"],"URL":"https:\/\/doi.org\/10.1186\/s40659-026-00680-3","relation":{},"ISSN":["0717-6287"],"issn-type":[{"value":"0717-6287","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,8]]},"assertion":[{"value":"5 October 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 February 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 June 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare that there are no financial, personal, or professional conflicts of interest that could be perceived to have influenced the work reported in this manuscript. No author has received direct payment, holds financial interests, or maintains affiliations with organizations that could be viewed as having a vested interest in the subject matter. All authors affirm the integrity and impartiality of the research and its conclusions.","order":3,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"55"}}