{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,19]],"date-time":"2026-10-19T15:40:58Z","timestamp":1792424458034,"version":"4.5.0"},"reference-count":93,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,7,22]],"date-time":"2026-07-22T00:00:00Z","timestamp":1784678400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,10,19]],"date-time":"2026-10-19T00:00:00Z","timestamp":1792368000000},"content-version":"vor","delay-in-days":89,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/501100001411","name":"Indian Council of Medical Research","doi-asserted-by":"publisher","award":["5\/10\/FR\/57\/2020-RBMCH"],"award-info":[{"award-number":["5\/10\/FR\/57\/2020-RBMCH"]}],"id":[{"id":"10.13039\/501100001411","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100007338","name":"All-India Institute of Medical Sciences","doi-asserted-by":"publisher","award":["A-1032"],"award-info":[{"award-number":["A-1032"]}],"id":[{"id":"10.13039\/501100007338","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Cell Commun Signal"],"DOI":"10.1186\/s12964-026-03069-7","type":"journal-article","created":{"date-parts":[[2026,7,22]],"date-time":"2026-07-22T09:24:39Z","timestamp":1784712279000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Reduced insulin\/IGF-1 signaling extends reproductive span through somatic gonadal collagen-mediated maintenance of oocyte and embryo quality in Caenorhabditis elegans"],"prefix":"10.1186","volume":"24","author":[{"given":"Neha","family":"Kaushik","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Soumya","family":"Rastogi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vaibhav","family":"Jain","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sonia","family":"Verma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Deepak","family":"Pandey","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mona","family":"Sharma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ashutosh","family":"Halder","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simon","family":"Gregory","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0906-4559","authenticated-orcid":false,"given":"Neeraj","family":"Kumar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,7,22]]},"reference":[{"key":"3069_CR1","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1093\/humupd\/8.2.141","volume":"8","author":"ER Te Velde","year":"2002","unstructured":"Te Velde ER, Pearson PL. The Variability of Female Reproductive Ageing. Hum Reprod Update. 2002;8:141\u201354.","journal-title":"Hum Reprod Update"},{"key":"3069_CR2","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1016\/S0015-0282(99)00231-9","volume":"72","author":"P Goud","year":"1999","unstructured":"Goud P, Goud A, Van Oostveldt P, Van der Elst J, Dhont M. Fertilization Abnormalities and Pronucleus Size Asynchrony after Intracytoplasmic Sperm Injection Are Related to Oocyte Postmaturity. Fertil Steril. 1999;72:245\u201352.","journal-title":"Fertil Steril"},{"key":"3069_CR3","doi-asserted-by":"publisher","first-page":"104804","DOI":"10.1016\/j.rbmo.2025.104804","volume":"50","author":"RA Anderson","year":"2025","unstructured":"Anderson RA, Marston AL, Telfer EE. Oocyte Development: It\u2019s All about Quality. Reprod Biomed Online. 2025;50:104804.","journal-title":"Reprod Biomed Online"},{"key":"3069_CR4","doi-asserted-by":"publisher","first-page":"1335","DOI":"10.3390\/antiox11071335","volume":"11","author":"V Immediata","year":"2022","unstructured":"Immediata V, Ronchetti C, Spadaro D, Cirillo F, Levi-Setti P.E. Oxidative Stress and Human Ovarian Response\u2014from Somatic Ovarian Cells to Oocytes Damage: A Clinical Comprehensive Narrative Review. Antioxidants. 2022;11:1335.","journal-title":"Antioxidants"},{"key":"3069_CR5","doi-asserted-by":"publisher","first-page":"e58619","DOI":"10.7554\/eLife.58619","volume":"9","author":"TA Starich","year":"2020","unstructured":"Starich TA, Bai X, Greenstein D. Gap Junctions Deliver Malonyl-CoA from Soma to Germline to Support Embryogenesis in Caenorhabditis elegans. Elife. 2020;9:e58619.","journal-title":"Elife"},{"key":"3069_CR6","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1002\/rmb2.12292","volume":"19","author":"MH Alam","year":"2020","unstructured":"Alam MH, Miyano T. Interaction between Growing Oocytes and Granulosa Cells in Vitro. Reprod Med Biol. 2020;19:13\u201323.","journal-title":"Reprod Med Biol"},{"key":"3069_CR7","doi-asserted-by":"publisher","first-page":"6","DOI":"10.3390\/jdb7010006","volume":"7","author":"CA Kelley","year":"2019","unstructured":"Kelley CA, Cram EJ. Regulation of Actin Dynamics in the C. elegans Somatic Gonad. J Dev Biol. 2019;7:6.","journal-title":"J Dev Biol"},{"key":"3069_CR8","doi-asserted-by":"publisher","first-page":"e1000295","DOI":"10.1371\/journal.pgen.1000295","volume":"4","author":"S Andux","year":"2008","unstructured":"Andux S, Ellis RE. Apoptosis Maintains Oocyte Quality in Aging Caenorhabditis elegans Females. PLoS Genet. 2008;4:e1000295.","journal-title":"PLoS Genet"},{"key":"3069_CR9","doi-asserted-by":"publisher","first-page":"e25","DOI":"10.1371\/journal.pgen.0030025","volume":"3","author":"SE Hughes","year":"2007","unstructured":"Hughes SE, Evason K, Xiong C, Kornfeld K. Genetic and Pharmacological Factors That Influence Reproductive Aging in Nematodes. PLoS Genet. 2007;3:e25.","journal-title":"PLoS Genet"},{"key":"3069_CR10","doi-asserted-by":"publisher","first-page":"299","DOI":"10.1016\/j.cell.2010.09.013","volume":"143","author":"S Luo","year":"2010","unstructured":"Luo S, Kleemann GA, Ashraf JM, Shaw WM, Murphy CT. TGF-\u03b2 and Insulin Signaling Regulate Reproductive Aging via Oocyte and Germline Quality Maintenance. Cell. 2010;143:299\u2013312.","journal-title":"Cell"},{"key":"3069_CR11","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1006\/dbio.1996.8429","volume":"181","author":"J McCarter","year":"1997","unstructured":"McCarter J, Bartlett B, Dang T, Schedl T. Soma\u2013Germ Cell Interactions in Caenorhabditis elegans: Multiple Events of Hermaphrodite Germline Development Require the Somatic Sheath and Spermathecal Lineages. Dev Biol. 1997;181:121\u201343.","journal-title":"Dev Biol"},{"key":"3069_CR12","doi-asserted-by":"publisher","first-page":"1009","DOI":"10.1016\/S0960-9822(99)80449-7","volume":"9","author":"B Kraemer","year":"1999","unstructured":"Kraemer B, Crittenden S, Gallegos M, Moulder G, Barstead R, Kimble J, Wickens M. NANOS-3 and FBF Proteins Physically Interact to Control the Sperm\u2013Oocyte Switch in Caenorhabditis elegans. Curr Biol. 1999;9:1009\u201318.","journal-title":"Curr Biol"},{"key":"3069_CR13","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1016\/0092-8674(87)90128-0","volume":"51","author":"J Austin","year":"1987","unstructured":"Austin J, Kimble J. Glp-1 Is Required in the Germ Line for Regulation of the Decision between Mitosis and Meiosis in C. elegans. Cell. 1987;51:589\u201399.","journal-title":"Cell"},{"key":"3069_CR14","doi-asserted-by":"publisher","first-page":"5332","DOI":"10.1073\/pnas.0711911105","volume":"105","author":"MA Cant","year":"2008","unstructured":"Cant MA, Johnstone RA. Reproductive Conflict and the Separation of Reproductive Generations in Humans. Proc Natl Acad Sci. 2008;105:5332\u20136.","journal-title":"Proc Natl Acad Sci"},{"key":"3069_CR15","doi-asserted-by":"publisher","first-page":"e1000789","DOI":"10.1371\/journal.pgen.1000789","volume":"5","author":"S Luo","year":"2009","unstructured":"Luo S, Shaw WM, Ashraf J, Murphy CT. TGF-\u00df Sma\/Mab Signaling Mutations Uncouple Reproductive Aging from Somatic Aging. PLoS Genet. 2009;5:e1000789.","journal-title":"PLoS Genet"},{"key":"3069_CR16","doi-asserted-by":"publisher","first-page":"8084","DOI":"10.1073\/pnas.0400848101","volume":"101","author":"C Huang","year":"2004","unstructured":"Huang C, Xiong C, Kornfeld K. Measurements of Age-Related Changes of Physiological Processes That Predict Lifespan of Caenorhabditis elegans. Proc Natl Acad Sci. 2004;101:8084\u20139.","journal-title":"Proc Natl Acad Sci"},{"key":"3069_CR17","doi-asserted-by":"publisher","first-page":"703","DOI":"10.1093\/genetics\/148.2.703","volume":"148","author":"HA Tissenbaum","year":"1998","unstructured":"Tissenbaum HA, Ruvkun G. An Insulin-like Signaling Pathway Affects Both Longevity and Reproduction in Caenorhabditis elegans. Genetics. 1998;148:703\u201317.","journal-title":"Genetics"},{"key":"3069_CR18","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1534\/genetics.111.130450","volume":"189","author":"SE Hughes","year":"2011","unstructured":"Hughes SE, Huang C, Kornfeld K. Identification of Mutations That Delay Somatic or Reproductive Aging of Caenorhabditis elegans. Genetics. 2011;189:341\u201356.","journal-title":"Genetics"},{"key":"3069_CR19","doi-asserted-by":"publisher","first-page":"753","DOI":"10.1016\/j.cub.2018.01.052","volume":"28","author":"NM Templeman","year":"2018","unstructured":"Templeman NM, Luo S, Kaletsky R, Shi C, Ashraf J, Keyes W, Murphy CT. Insulin Signaling Regulates Oocyte Quality Maintenance with Age via Cathepsin B Activity. Curr Biol. 2018;28:753\u201360.","journal-title":"Curr Biol"},{"key":"3069_CR20","doi-asserted-by":"publisher","first-page":"1635","DOI":"10.1016\/j.cub.2007.08.058","volume":"17","author":"WM Shaw","year":"2007","unstructured":"Shaw WM, Luo S, Landis J, Ashraf J, Murphy CT. The C. elegans TGF-\u03b2 Dauer Pathway Regulates Longevity via Insulin Signaling. Curr Biol. 2007;17:1635\u201345.","journal-title":"Curr Biol"},{"key":"3069_CR21","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1006\/dbio.1999.9356","volume":"212","author":"DH Hall","year":"1999","unstructured":"Hall DH, Winfrey VP, Blaeuer G, Hoffman LH, Furuta T, Rose KL, Hobert O, Greenstein D. Ultrastructural Features of the Adult Hermaphrodite Gonad of Caenorhabditis elegans: Relations between the Germ Line and Soma. Dev Biol. 1999;212:101\u201323.","journal-title":"Dev Biol"},{"key":"3069_CR22","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1146\/annurev.physiol.61.1.283","volume":"61","author":"TW White","year":"1999","unstructured":"White TW, Paul DL. Genetic Diseases and Gene Knockouts Reveal Diverse Connexin Functions. Annu Rev Physiol. 1999;61:283\u2013310.","journal-title":"Annu Rev Physiol"},{"key":"3069_CR23","doi-asserted-by":"publisher","first-page":"680","DOI":"10.1083\/jcb.71.2.680","volume":"71","author":"E Anderson","year":"1976","unstructured":"Anderson E, Albertini DF. Gap Junctions between the Oocyte and Companion Follicle Cells in the Mammalian Ovary. J Cell Biol. 1976;71:680\u20136.","journal-title":"J Cell Biol"},{"key":"3069_CR24","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1006\/dbio.1996.8429","volume":"181","author":"J McCarter","year":"1997","unstructured":"McCarter J, Bartlett B, Dang T, Schedl T. Soma\u2013Germ Cell Interactions in Caenorhabditis elegans: Multiple Events of Hermaphrodite Germline Development Require the Somatic Sheath and Spermathecal Lineages. Dev Biol. 1997;181:121\u201343.","journal-title":"Dev Biol"},{"key":"3069_CR25","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1006\/dbio.1998.9109","volume":"205","author":"J McCarter","year":"1999","unstructured":"McCarter J, Bartlett B, Dang T, Schedl T. On the Control of Oocyte Meiotic Maturation and Ovulation In Caenorhabditis elegans. Dev Biol. 1999;205:111\u201328.","journal-title":"Dev Biol"},{"key":"3069_CR26","doi-asserted-by":"publisher","unstructured":"Govindan JA, Nadarajan S, Kim S, Starich TA, Greenstein D. Somatic cAMP Signaling Regulates MSP-Dependent Oocyte Growth and Meiotic Maturation in C. elegans. 2009;136(13):2211\u201321. https:\/\/doi.org\/10.1242\/dev.034595.","DOI":"10.1242\/dev.034595"},{"key":"3069_CR27","doi-asserted-by":"publisher","first-page":"1223","DOI":"10.1007\/s43032-019-00137-x","volume":"27","author":"A Arroyo","year":"2020","unstructured":"Arroyo A, Kim B, Yeh J. Luteinizing Hormone Action in Human Oocyte Maturation and Quality: Signaling Pathways, Regulation, and Clinical Impact. Reprod Sci. 2020;27:1223\u201352.","journal-title":"Reprod Sci"},{"key":"3069_CR28","doi-asserted-by":"publisher","first-page":"54","DOI":"10.3390\/ijms23010054","volume":"23","author":"E Nagyov\u00e1","year":"2021","unstructured":"Nagyov\u00e1 E, N\u011bmcov\u00e1 L, Camaioni A. Cumulus Extracellular Matrix Is an Important Part of Oocyte Microenvironment in Ovarian Follicles: Its Remodeling and Proteolytic Degradation. Int J Mol Sci. 2021;23:54.","journal-title":"Int J Mol Sci"},{"key":"3069_CR29","doi-asserted-by":"publisher","first-page":"110006","DOI":"10.1016\/j.cellsig.2021.110006","volume":"84","author":"A Amran","year":"2021","unstructured":"Amran A, Pigatto L, Pocock R, Gopal S. Functions of the Extracellular Matrix in Development: Lessons from Caenorhabditis elegans. Cell Signal. 2021;84:110006.","journal-title":"Cell Signal"},{"key":"3069_CR30","doi-asserted-by":"publisher","first-page":"4200","DOI":"10.1038\/s41467-024-48283-4","volume":"15","author":"A Amran","year":"2024","unstructured":"Amran A, Pigatto L, Farley J, Godini R, Pocock R, Gopal S. The Matrisome Landscape Controlling in Vivo Germ Cell Fates. Nat Commun. 2024;15:4200.","journal-title":"Nat Commun"},{"key":"3069_CR31","doi-asserted-by":"publisher","first-page":"106667","DOI":"10.1016\/j.jsbmb.2024.106667","volume":"247","author":"H Xi","year":"2025","unstructured":"Xi H, Chen X, Wang X, Jiang F, Niu D. Role of Programmed Cell Death in Mammalian Ovarian Follicular Atresia. J Steroid Biochem Mol Biol. 2025;247:106667.","journal-title":"J Steroid Biochem Mol Biol"},{"key":"3069_CR32","doi-asserted-by":"publisher","first-page":"39082","DOI":"10.18632\/oncotarget.9681","volume":"7","author":"Y de la Guardia","year":"2016","unstructured":"de la Guardia Y, Gilliat AF, Hellberg J, Rennert P, Cabreiro F, Gems D. Run-on of Germline Apoptosis Promotes Gonad Senescence in C. elegans. Oncotarget. 2016;7:39082.","journal-title":"Oncotarget"},{"key":"3069_CR33","doi-asserted-by":"publisher","first-page":"8084","DOI":"10.1073\/pnas.0400848101","volume":"101","author":"C Huang","year":"2004","unstructured":"Huang C, Xiong C, Kornfeld K. Measurements of Age-Related Changes of Physiological Processes That Predict Lifespan of Caenorhabditis elegans. Proc Natl Acad Sci. 2004;101:8084\u20139.","journal-title":"Proc Natl Acad Sci"},{"key":"3069_CR34","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1046\/j.1474-9728.2002.00016.x","volume":"1","author":"M Tu","year":"2002","unstructured":"Tu M, Yin C, Tatar M. Impaired Ovarian Ecdysone Synthesis of Drosophila Melanogaster Insulin Receptor Mutants. Aging Cell. 2002;1:158\u201360.","journal-title":"Aging Cell"},{"key":"3069_CR35","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1126\/science.1086336","volume":"301","author":"DH Castrillon","year":"2003","unstructured":"Castrillon DH, Miao L, Kollipara R, Horner JW, DePinho RA. Suppression of Ovarian Follicle Activation in Mice by the Transcription Factor Foxo3a. Science. 2003;301:215\u20138.","journal-title":"Science"},{"key":"3069_CR36","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1038\/35030105","volume":"407","author":"DJ Burks","year":"2000","unstructured":"Burks DJ, de Mora JF, Schubert M, Withers DJ, Myers MG, Towery HH, Altamuro SL, Flint CL, White MF. IRS-2 Pathways Integrate Female Reproduction and Energy Homeostasis. Nature. 2000;407:377.","journal-title":"Nature"},{"key":"3069_CR37","doi-asserted-by":"publisher","first-page":"1518","DOI":"10.1016\/j.fertnstert.2006.03.054","volume":"86","author":"WJ Watkins","year":"2006","unstructured":"Watkins WJ, Umbers AJ, Woad KJ, Harris SE, Winship IM, Gersak K, Shelling AN. Mutational Screening of FOXO3A and FOXO1A in Women with Premature Ovarian Failure. Fertil Steril. 2006;86:1518\u201321.","journal-title":"Fertil Steril"},{"key":"3069_CR38","doi-asserted-by":"publisher","first-page":"216","DOI":"10.1093\/humrep\/dem255","volume":"23","author":"TD Gallardo","year":"2007","unstructured":"Gallardo TD, John GB, Bradshaw K, Welt C, Reijo-Pera R, Vogt PH, Touraine P, Bione S, Toniolo D, Nelson LM. Sequence Variation at the Human FOXO3 Locus: A Study of Premature Ovarian Failure and Primary Amenorrhea. Hum Reprod. 2007;23:216\u201321.","journal-title":"Hum Reprod"},{"key":"3069_CR39","doi-asserted-by":"publisher","first-page":"444","DOI":"10.1002\/mrd.22806","volume":"84","author":"D Das","year":"2017","unstructured":"Das D, Arur S. Conserved Insulin Signaling in the Regulation of Oocyte Growth, Development, and Maturation. Mol Reprod Dev. 2017;84:444\u201359.","journal-title":"Mol Reprod Dev"},{"key":"3069_CR40","doi-asserted-by":"publisher","first-page":"694","DOI":"10.1093\/molehr\/6.8.694","volume":"6","author":"H Louhio","year":"2000","unstructured":"Louhio H, Hovatta O, Sj\u00f6berg J, Tuuri T. The Effects of Insulin, and Insulin-like Growth Factors I and II on Human Ovarian Follicles in Long-Term Culture. Mol Hum Reprod. 2000;6:694\u20138.","journal-title":"Mol Hum Reprod"},{"key":"3069_CR41","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1016\/j.devcel.2013.09.008","volume":"27","author":"AL Lopez","year":"2013","unstructured":"Lopez AL, Chen J, Joo H-J, Drake M, Shidate M, Kseib C, Arur S. DAF-2 and ERK Couple Nutrient Availability to Meiotic Progression during Caenorhabditis elegans Oogenesis. Dev Cell. 2013;27:227\u201340.","journal-title":"Dev Cell"},{"key":"3069_CR42","doi-asserted-by":"publisher","first-page":"671","DOI":"10.1242\/dev.042523","volume":"137","author":"D Michaelson","year":"2010","unstructured":"Michaelson D, Korta DZ, Capua Y, Hubbard EJ. A. Insulin Signaling Promotes Germline Proliferation in C. elegans. Development. 2010;137:671\u201380.","journal-title":"Development"},{"key":"3069_CR43","doi-asserted-by":"publisher","first-page":"529","DOI":"10.1186\/s12964-025-02510-7","volume":"23","author":"N Kaushik","year":"2025","unstructured":"Kaushik N, Rastogi S, Kapadia S, Jain V, Verma S, Pandey D, Sharma M, Gupta S, Halder A, Gregory S. Temporal Transcriptomic Profiling Reveals Distinct Age-Associated Gene Expression Signatures in Gonads under Reduced Insulin\/IGF-1 Signaling in Caenorhabditis elegans. Cell Commun Signal. 2025;23:529.","journal-title":"Cell Commun Signal"},{"key":"3069_CR44","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13059-014-0550-8","volume":"15","author":"MI Love","year":"2014","unstructured":"Love MI, Huber W, Anders S. Moderated Estimation of Fold Change and Dispersion for RNA-Seq Data with DESeq2. Genome Biol. 2014;15:1\u201321.","journal-title":"Genome Biol"},{"key":"3069_CR45","doi-asserted-by":"publisher","first-page":"D638","DOI":"10.1093\/nar\/gkac1000","volume":"51","author":"D Szklarczyk","year":"2023","unstructured":"Szklarczyk D, Kirsch R, Koutrouli M, Nastou K, Mehryary F, Hachilif R, Gable AL, Fang T, Doncheva NT, Pyysalo S. The STRING Database in 2023: Protein\u2013Protein Association Networks and Functional Enrichment Analyses for Any Sequenced Genome of Interest. Nucleic Acids Res. 2023;51:D638\u201346.","journal-title":"Nucleic Acids Res"},{"key":"3069_CR46","doi-asserted-by":"publisher","first-page":"2498","DOI":"10.1101\/gr.1239303","volume":"13","author":"P Shannon","year":"2003","unstructured":"Shannon P, Markiel A, Ozier O, Baliga NS, Wang JT, Ramage D, Amin N, Schwikowski B, Ideker T. Cytoscape: A Software Environment for Integrated Models of Biomolecular Interaction Networks. Genome Res. 2003;13:2498\u2013504.","journal-title":"Genome Res"},{"key":"3069_CR47","unstructured":"Oliveros JC. Venny. An Interactive Tool for Comparing Lists with Venn\u2019s Diagrams. 2007\/2015\/2024. http:\/\/bioinfogp.cnb.csic.es\/tools\/venny\/index.html.\u00a0accessed March 23, 2025."},{"key":"3069_CR48","doi-asserted-by":"publisher","first-page":"1805","DOI":"10.1093\/bioinformatics\/bts251","volume":"28","author":"X Jiao","year":"2012","unstructured":"Jiao X, Sherman BT, Huang DW, Stephens R, Baseler MW, Lane HC, Lempicki RA. DAVID-WS: A Stateful Web Service to Facilitate Gene\/Protein List Analysis. Bioinformatics. 2012;28:1805\u20136.","journal-title":"Bioinformatics"},{"key":"3069_CR49","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1016\/S1046-2023(03)00050-1","volume":"30","author":"RS Kamath","year":"2003","unstructured":"Kamath RS, Ahringer J. Genome-Wide RNAi Screening in Caenorhabditis elegans. Methods. 2003;30:313\u201321.","journal-title":"Methods"},{"key":"3069_CR50","doi-asserted-by":"publisher","first-page":"a004978","DOI":"10.1101\/cshperspect.a004978","volume":"3","author":"S Ricard-Blum","year":"2011","unstructured":"Ricard-Blum S. The Collagen Family. Cold Spring Harb Perspect Biol. 2011;3:a004978.","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"3069_CR51","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12864-019-5893-9","volume":"20","author":"M Han","year":"2019","unstructured":"Han M, Wei G, McManus CE, Hillier LW, Reinke V. Isolated C. elegans Germ Nuclei Exhibit Distinct Genomic Profiles of Histone Modification and Gene Expression. BMC Genomics. 2019;20:1\u201315.","journal-title":"BMC Genomics"},{"key":"3069_CR52","doi-asserted-by":"publisher","first-page":"e30201","DOI":"10.7554\/eLife.30201","volume":"6","author":"C-YS Lee","year":"2017","unstructured":"Lee C-YS, Lu T, Seydoux G. Nanos Promotes Epigenetic Reprograming of the Germline by Down-Regulation of the THAP Transcription Factor LIN-15B. Elife. 2017;6:e30201.","journal-title":"Elife"},{"key":"3069_CR53","doi-asserted-by":"publisher","first-page":"1765","DOI":"10.1534\/g3.114.012351","volume":"4","author":"MA Ortiz","year":"2014","unstructured":"Ortiz MA, Noble D, Sorokin EP, Kimble J. A New Dataset of Spermatogenic vs. Oogenic Transcriptomes in the Nematode Caenorhabditis elegans. G3 Genes Genomes Genet. 2014;4:1765\u201372.","journal-title":"G3 Genes Genomes Genet"},{"key":"3069_CR54","doi-asserted-by":"publisher","first-page":"491","DOI":"10.1038\/ncb2720","volume":"15","author":"CG Riedel","year":"2013","unstructured":"Riedel CG, Dowen RH, Lourenco GF, Kirienko NV, Heimbucher T, West JA, Bowman SK, Kingston RE, Dillin A, Asara JM. DAF-16 Employs the Chromatin Remodeller SWI\/SNF to Promote Stress Resistance and Longevity. Nat Cell Biol. 2013;15:491\u2013501.","journal-title":"Nat Cell Biol"},{"key":"3069_CR55","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1038\/nature01789","volume":"424","author":"CT Murphy","year":"2003","unstructured":"Murphy CT, McCarroll SA, Bargmann CI, Fraser A, Kamath RS, Ahringer J, Li H, Kenyon C. Genes That Act Downstream of DAF-16 to Influence the Lifespan of Caenorhabditis elegans. Nature. 2003;424:277\u201383.","journal-title":"Nature"},{"key":"3069_CR56","doi-asserted-by":"publisher","first-page":"12462","DOI":"10.3390\/ijms222212462","volume":"22","author":"N Kaushik","year":"2021","unstructured":"Kaushik N, Rastogi S, Verma S, Pandey D, Halder A, Mukhopadhyay A, Kumar N. Transcriptome Analysis of Insulin Signaling-Associated Transcription Factors in C. elegans Reveal Their Genome-Wide Target Genes Specificity and Complexity. Int J Mol Sci. 2021;22:12462.","journal-title":"Int J Mol Sci"},{"key":"3069_CR57","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1046\/j.1474-9728.2003.00043.x","volume":"2","author":"J McElwee","year":"2003","unstructured":"McElwee J, Bubb K, Thomas JH. Transcriptional Outputs of the Caenorhabditis elegans Forkhead Protein DAF-16. Aging Cell. 2003;2:111\u201321.","journal-title":"Aging Cell"},{"key":"3069_CR58","doi-asserted-by":"publisher","first-page":"41418","DOI":"10.18632\/oncotarget.6282","volume":"6","author":"N Kumar","year":"2015","unstructured":"Kumar N, Jain V, Singh A, Jagtap U, Verma S, Mukhopadhyay A. Genome-Wide Endogenous DAF-16\/FOXO Recruitment Dynamics during Lowered Insulin Signalling in C. elegans. Oncotarget. 2015;6:41418.","journal-title":"Oncotarget"},{"key":"3069_CR59","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1038\/nature14021","volume":"519","author":"CY Ewald","year":"2015","unstructured":"Ewald CY, Landis JN, Abate JP, Murphy CT, Blackwell TK. Dauer-Independent Insulin\/IGF-1-Signalling Implicates Collagen Remodelling in Longevity. Nature. 2015;519:97\u2013101.","journal-title":"Nature"},{"key":"3069_CR60","doi-asserted-by":"publisher","first-page":"524","DOI":"10.1111\/j.1474-9726.2009.00501.x","volume":"8","author":"RP Oliveira","year":"2009","unstructured":"Oliveira RP, Abate JP, Dilks K, Landis J, Ashraf J, Murphy CT, Blackwell TK. Condition-adapted Stress and Longevity Gene Regulation by Caenorhabditis elegans SKN\u20101\/Nrf. Aging Cell. 2009;8:524\u201341.","journal-title":"Aging Cell"},{"key":"3069_CR61","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1186\/s12864-016-2837-5","volume":"17","author":"J Brunquell","year":"2016","unstructured":"Brunquell J, Morris S, Lu Y, Cheng F, Westerheide SD. The Genome-Wide Role of HSF-1 in the Regulation of Gene Expression in Caenorhabditis elegans. BMC Genomics. 2016;17:559.","journal-title":"BMC Genomics"},{"key":"3069_CR62","doi-asserted-by":"publisher","first-page":"1679","DOI":"10.1534\/g3.119.400044","volume":"9","author":"S Sural","year":"2019","unstructured":"Sural S, Lu T-C, Jung SA, Hsu A-L. HSB-1 Inhibition and HSF-1 Overexpression Trigger Overlapping Transcriptional Changes to Promote Longevity in Caenorhabditis elegans. G3 Genes Genomes Genet. 2019;9:1679\u201392.","journal-title":"G3 Genes Genomes Genet"},{"key":"3069_CR63","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1002\/evl3.108","volume":"3","author":"MI Lind","year":"2019","unstructured":"Lind MI, Ravindran S, Sekajova Z, Carlsson H, Hinas A, Maklakov AA. Experimentally Reduced Insulin\/IGF-1 Signaling in Adulthood Extends Lifespan of Parents and Improves Darwinian Fitness of Their Offspring. Evol Lett. 2019;3:207\u201316.","journal-title":"Evol Lett"},{"key":"3069_CR64","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1038\/20694","volume":"399","author":"H Hsin","year":"1999","unstructured":"Hsin H, Kenyon C. Signals from the Reproductive System Regulate the Lifespan of C. elegans. Nature. 1999;399:362\u20136.","journal-title":"Nature"},{"key":"3069_CR65","doi-asserted-by":"publisher","first-page":"6368","DOI":"10.1073\/pnas.0709128105","volume":"105","author":"T Flatt","year":"2008","unstructured":"Flatt T, Min K-J, D\u2019Alterio C, Villa-Cuesta E, Cumbers J, Lehmann R, Jones DL, Tatar M. Drosophila Germ-Line Modulation of Insulin Signaling and Lifespan. Proc Natl Acad Sci. 2008;105:6368\u201373.","journal-title":"Proc Natl Acad Sci"},{"key":"3069_CR66","doi-asserted-by":"publisher","first-page":"e1000639","DOI":"10.1371\/journal.pgen.1000639","volume":"5","author":"A Ghazi","year":"2009","unstructured":"Ghazi A, Henis-Korenblit S, Kenyon CA. Transcription Elongation Factor That Links Signals from the Reproductive System to Lifespan Extension in Caenorhabditis elegans. PLoS Genet. 2009;5:e1000639.","journal-title":"PLoS Genet"},{"key":"3069_CR67","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.matbio.2026.02.002","volume":"145","author":"V Stolzenbach","year":"2026","unstructured":"Stolzenbach V, Griffin SS, Goncalves L, Agarwal P, Zaidel-Bar R, Cram EJ. A Switch in Collagen Expression Regulates Cessation of Distal Tip Cell Migration in C. elegans. Matrix Biol. 2026;145:1\u201314.","journal-title":"Matrix Biol"},{"key":"3069_CR68","doi-asserted-by":"publisher","first-page":"1379","DOI":"10.1534\/genetics.111.133173","volume":"190","author":"Y Kubota","year":"2012","unstructured":"Kubota Y, Nagata K, Sugimoto A, Nishiwaki K. Tissue Architecture in the Caenorhabditis elegans Gonad Depends on Interactions among Fibulin-1, Type IV Collagen and the ADAMTS Extracellular Protease. Genetics. 2012;190:1379\u201388.","journal-title":"Genetics"},{"key":"3069_CR69","doi-asserted-by":"publisher","first-page":"1171","DOI":"10.1083\/jcb.137.5.1171","volume":"137","author":"PL Graham","year":"1997","unstructured":"Graham PL, Johnson JJ, Wang S, Sibley MH, Gupta MC, Kramer JM. Type IV Collagen Is Detectable in Most, but Not All, Basement Membranes of Caenorhabditis elegans and Assembles on Tissues That Do Not Express It. J Cell Biol. 1997;137:1171\u201383.","journal-title":"J Cell Biol"},{"key":"3069_CR70","doi-asserted-by":"publisher","first-page":"1186","DOI":"10.3390\/genes12081186","volume":"12","author":"WS Parkes","year":"2021","unstructured":"Parkes WS, Amargant F, Zhou LT, Villanueva CE, Duncan FE, Pritchard MT. Hyaluronan and Collagen Are Prominent Extracellular Matrix Components in Bovine and Porcine Ovaries. Genes. 2021;12:1186.","journal-title":"Genes"},{"key":"3069_CR71","doi-asserted-by":"publisher","first-page":"718522","DOI":"10.3389\/fcell.2021.718522","volume":"9","author":"A Scharf","year":"2021","unstructured":"Scharf A, Pohl F, Egan BM, Kocsisova Z, Kornfeld K. Reproductive Aging in Caenorhabditis elegans: From Molecules to Ecology. Front Cell Dev Biol. 2021;9:718522.","journal-title":"Front Cell Dev Biol"},{"key":"3069_CR72","doi-asserted-by":"publisher","first-page":"C1201","DOI":"10.1152\/ajpcell.00195.2023","volume":"325","author":"H Reich","year":"2023","unstructured":"Reich H, Savage-Dunn C. Signaling Circuits and the Apical Extracellular Matrix in Aging: Connections Identified in the Nematode Caenorhabditis elegans. Am J Physiol -Cell Physiol. 2023;325:C1201\u201311.","journal-title":"Am J Physiol -Cell Physiol"},{"key":"3069_CR73","doi-asserted-by":"publisher","unstructured":"Page AP, Johnstone IL. The cuticle. WormBook. 2007;19:1\u201315. https:\/\/doi.org\/10.1895\/wormbook.1.138.1.","DOI":"10.1895\/wormbook.1.138.1"},{"key":"3069_CR74","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1002\/bies.950160307","volume":"16","author":"IL Johnstone","year":"1994","unstructured":"Johnstone IL. The Cuticle of the Nematode Caenorhabditis elegans: A Complex Collagen Structure. BioEssays. 1994;16:171\u20138.","journal-title":"BioEssays"},{"key":"3069_CR75","doi-asserted-by":"publisher","first-page":"110006","DOI":"10.1016\/j.cellsig.2021.110006","volume":"84","author":"A Amran","year":"2021","unstructured":"Amran A, Pigatto L, Pocock R, Gopal S. Functions of the Extracellular Matrix in Development: Lessons from Caenorhabditis elegans. Cell Signal. 2021;84:110006.","journal-title":"Cell Signal"},{"key":"3069_CR76","doi-asserted-by":"publisher","first-page":"276","DOI":"10.1038\/s41467-023-44409-2","volume":"15","author":"AC Teuscher","year":"2024","unstructured":"Teuscher AC, Statzer C, Goyala A, Domenig SA, Schoen I, Hess M, Hofer AM, Fossati A, Vogel V, Goksel O. Longevity Interventions Modulate Mechanotransduction and Extracellular Matrix Homeostasis in C. elegans. Nat Commun. 2024;15:276.","journal-title":"Nat Commun"},{"key":"3069_CR77","doi-asserted-by":"publisher","first-page":"54","DOI":"10.3390\/ijms23010054","volume":"23","author":"E Nagyov\u00e1","year":"2021","unstructured":"Nagyov\u00e1 E, N\u011bmcov\u00e1 L, Camaioni A. Cumulus Extracellular Matrix Is an Important Part of Oocyte Microenvironment in Ovarian Follicles: Its Remodeling and Proteolytic Degradation. Int J Mol Sci. 2021;23:54.","journal-title":"Int J Mol Sci"},{"key":"3069_CR78","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1038\/nature14021","volume":"519","author":"CY Ewald","year":"2015","unstructured":"Ewald CY, Landis JN, Abate JP, Murphy CT, Blackwell TK. Dauer-Independent Insulin\/IGF-1-Signalling Implicates Collagen Remodelling in Longevity. Nature. 2015;519:97\u2013101.","journal-title":"Nature"},{"key":"3069_CR79","doi-asserted-by":"publisher","first-page":"e12896","DOI":"10.1111\/acel.12896","volume":"18","author":"S Li","year":"2019","unstructured":"Li S, Zhao H, Zhang P, Liang C, Zhang Y, Hsu A, Dong M. DAF-16 Stabilizes the Aging Transcriptome and Is Activated in Mid\u2010aged Caenorhabditis elegans to Cope with Internal Stress. Aging Cell. 2019;18:e12896.","journal-title":"Aging Cell"},{"key":"3069_CR80","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1038\/s41514-025-00216-1","volume":"11","author":"G Wang","year":"2025","unstructured":"Wang G, Yang R, Zhang H. Ovarian Vascular Aging: A Hidden Driver of Mid-Age Female Fertility Decline. Npj Aging. 2025;11:24.","journal-title":"Npj Aging"},{"key":"3069_CR81","doi-asserted-by":"publisher","first-page":"803","DOI":"10.1091\/mbc.4.8.803","volume":"4","author":"AD Levy","year":"1993","unstructured":"Levy AD, Yang J, Kramer JM. Molecular and Genetic Analyses of the Caenorhabditis elegans Dpy-2 and Dpy-10 Collagen Genes: A Variety of Molecular Alterations Affect Organismal Morphology. Mol Biol Cell. 1993;4:803\u201317.","journal-title":"Mol Biol Cell"},{"key":"3069_CR82","doi-asserted-by":"publisher","first-page":"e0243522","DOI":"10.1371\/journal.pone.0243522","volume":"16","author":"D Grushko","year":"2021","unstructured":"Grushko D, Boocholez H, Levine A, Cohen E. Temporal Requirements of SKN-1\/NRF as a Regulator of Lifespan and Proteostasis in Caenorhabditis elegans. PLoS ONE. 2021;16:e0243522.","journal-title":"PLoS ONE"},{"key":"3069_CR83","doi-asserted-by":"publisher","first-page":"1142","DOI":"10.1126\/science.1083701","volume":"300","author":"A-L Hsu","year":"2003","unstructured":"Hsu A-L, Murphy CT, Kenyon C. Regulation of Aging and Age-Related Disease by DAF-16 and Heat-Shock Factor. Science. 2003;300:1142\u20135.","journal-title":"Science"},{"key":"3069_CR84","doi-asserted-by":"publisher","first-page":"1191","DOI":"10.1111\/acel.12627","volume":"16","author":"JM Tullet","year":"2017","unstructured":"Tullet JM, Green JW, Au C, Benedetto A, Thompson MA, Clark E, Gilliat AF, Young A, Schmeisser K, Gems D. The SKN-1\/Nrf2 Transcription Factor Can Protect against Oxidative Stress and Increase Lifespan in C. elegans by Distinct Mechanisms. Aging Cell. 2017;16:1191\u20134.","journal-title":"Aging Cell"},{"key":"3069_CR85","doi-asserted-by":"publisher","unstructured":"Ragle JM, Pooranachithra M, Ashley GE, Cadena E, Blank B, Kang K, Chen C, Bhowmick AR, Mercado SH, Wells TE. Multiscale patterning of a model apical extracellular matrix revealed by systematic endogenous protein tagging. bioRxiv [Preprint]. 2025 May 14:2025.05.14.653803. https:\/\/doi.org\/10.1101\/2025.05.14.653803.","DOI":"10.1101\/2025.05.14.653803"},{"key":"3069_CR86","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1038\/s41536-022-00204-z","volume":"7","author":"M Loreti","year":"2022","unstructured":"Loreti M, Sacco A. The Jam Session between Muscle Stem Cells and the Extracellular Matrix in the Tissue Microenvironment. NPJ Regen Med. 2022;7:16.","journal-title":"NPJ Regen Med"},{"key":"3069_CR87","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1016\/j.ydbio.2005.11.026","volume":"290","author":"G Kao","year":"2006","unstructured":"Kao G, Huang C, Hedgecock EM, Hall DH, Wadsworth WG. The Role of the Laminin \u03b2 Subunit in Laminin Heterotrimer Assembly and Basement Membrane Function and Development in C. elegans. Dev Biol. 2006;290:211\u20139.","journal-title":"Dev Biol"},{"key":"3069_CR88","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1186\/s13048-025-01596-4","volume":"18","author":"Z Zhu","year":"2025","unstructured":"Zhu Z, Lei M, Guo R, Xu Y, Zhao Y, Wei C, Yang Q, Sun Y. Nicotinamide Riboside Supplementation Ameliorates Ovarian Dysfunction in a PCOS Mouse Model. J Ovarian Res. 2025;18:9.","journal-title":"J Ovarian Res"},{"key":"3069_CR89","doi-asserted-by":"publisher","first-page":"829","DOI":"10.1093\/molehr\/gap072","volume":"15","author":"MJ Prodoehl","year":"2009","unstructured":"Prodoehl MJ, Hatzirodos N, Irving-Rodgers HF, Zhao ZZ, Painter JN, Hickey TE, Gibson MA, Rainey WE, Carr BR, Mason HD. Genetic and Gene Expression Analyses of the Polycystic Ovary Syndrome Candidate Gene Fibrillin-3 and Other Fibrillin Family Members in Human Ovaries. Mol Hum Reprod. 2009;15:829\u201341.","journal-title":"Mol Hum Reprod"},{"key":"3069_CR90","doi-asserted-by":"publisher","first-page":"1476","DOI":"10.1080\/00016340601033741","volume":"85","author":"A-K Lind","year":"2006","unstructured":"Lind A-K, Weijdeg\u00e5rd B, Dahm-K\u00e4hler P, M\u00f6lne J, Sundfeldt K, Br\u00e4nnstr\u00f6m M. Collagens in the Human Ovary and Their Changes in the Perifollicular Stroma during Ovulation. Acta Obstet Gynecol Scand. 2006;85:1476\u201384.","journal-title":"Acta Obstet Gynecol Scand"},{"key":"3069_CR91","doi-asserted-by":"publisher","first-page":"8644","DOI":"10.1038\/s41598-019-45155-6","volume":"9","author":"M Griffiths","year":"2019","unstructured":"Griffiths M, Van Sinderen M, Rainczuk K, Dimitriadis E. miR-29c Overexpression and COL4A1 Downregulation in Infertile Human Endometrium Reduces Endometrial Epithelial Cell Adhesive Capacity in Vitro Implying Roles in Receptivity. Sci Rep. 2019;9:8644.","journal-title":"Sci Rep"},{"key":"3069_CR92","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1002\/mrd.23554","volume":"89","author":"H Du","year":"2022","unstructured":"Du H, Guo Y, Wu X, Gong Y. FOXL2 Regulates the Expression of the Col4a1 Collagen Gene in Chicken Granulosa Cells. Mol Reprod Dev. 2022;89:95\u2013103.","journal-title":"Mol Reprod Dev"},{"key":"3069_CR93","doi-asserted-by":"publisher","first-page":"3044","DOI":"10.3390\/ijms22063044","volume":"22","author":"NM Abogresha","year":"2021","unstructured":"Abogresha NM, Mohammed SS, Hosny MM, Abdallah HY, Gadallah AM, Greish SM. Diosmin Mitigates Cyclophosphamide Induced Premature Ovarian Insufficiency in Rat Model. Int J Mol Sci. 2021;22:3044.","journal-title":"Int J Mol Sci"}],"container-title":["Cell Communication and Signaling"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12964-026-03069-7","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12964-026-03069-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12964-026-03069-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,19]],"date-time":"2026-10-19T15:02:52Z","timestamp":1792422172000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12964-026-03069-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,22]]},"references-count":93,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["3069"],"URL":"https:\/\/doi.org\/10.1186\/s12964-026-03069-7","relation":{},"ISSN":["1478-811X"],"issn-type":[{"value":"1478-811X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,22]]},"assertion":[{"value":"24 September 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"10 July 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 July 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The study was approved by the Institute Ethics Committee of All India Institute of Medical Sciences (AIIMS), New Delhi (Ref No. IEC-120\/06.02.2020, dated 09\/03\/2020). Ethical approval was obtained as part of the institutional protocol, although specific consent to participate was not needed, as the research involved a model organism, which is exempt from human ethical review requirements.","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 no competing interests.","order":3,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"563"}}