{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,9]],"date-time":"2026-11-09T21:24:53Z","timestamp":1794259493771,"version":"4.9.0"},"reference-count":124,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,7,7]],"date-time":"2026-07-07T00:00:00Z","timestamp":1783382400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100013948","name":"Tempus Public Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100013948","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003827","name":"Hungarian Scientific Research Fund","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100003827","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Mechanisms of Ageing and Development"],"published-print":{"date-parts":[[2026,10]]},"DOI":"10.1016\/j.mad.2026.112225","type":"journal-article","created":{"date-parts":[[2026,7,8]],"date-time":"2026-07-08T15:31:54Z","timestamp":1783524714000},"page":"112225","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":2,"special_numbering":"C","title":["HSF1 in aging: The output selection model"],"prefix":"10.1016","volume":"233","author":[{"ORCID":"https:\/\/orcid.org\/0009-0007-3653-2729","authenticated-orcid":false,"given":"Saba","family":"Khatatneh","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4057-7678","authenticated-orcid":false,"given":"Csaba","family":"S\u0151ti","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6257-4849","authenticated-orcid":false,"given":"Mil\u00e1n","family":"Somogyv\u00e1ri","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"issue":"7","key":"10.1016\/j.mad.2026.112225_bib1","doi-asserted-by":"crossref","first-page":"1153","DOI":"10.1101\/gad.6.7.1153","article-title":"The human heat shock protein hsp70 interacts with HSF, the transcription factor that regulates heat shock gene expression","volume":"6","author":"Abravaya","year":"1992","journal-title":"Genes Dev."},{"key":"10.1016\/j.mad.2026.112225_bib2","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1038\/nrm2938","article-title":"Heat shock factors: integrators of cell stress, development and lifespan","volume":"11","author":"Akerfelt","year":"2010","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib3","doi-asserted-by":"crossref","first-page":"4949","DOI":"10.1128\/MCB.18.9.4949","article-title":"HSP90 interacts with and regulates the activity of heat shock factor 1 in Xenopus oocytes","volume":"18","author":"Ali","year":"1998","journal-title":"Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib4","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1007\/s10522-006-9053-7","article-title":"Protein homeostasis and molecular chaperones in aging","volume":"7","author":"Arslan","year":"2006","journal-title":"Biogerontology"},{"key":"10.1016\/j.mad.2026.112225_bib5","doi-asserted-by":"crossref","first-page":"360","DOI":"10.1126\/science.1253168","article-title":"HSF-1\u2013mediated cytoskeletal integrity determines thermotolerance and life span","volume":"346","author":"Baird","year":"2014","journal-title":"Science"},{"issue":"6","key":"10.1016\/j.mad.2026.112225_bib6","doi-asserted-by":"crossref","first-page":"1151","DOI":"10.1083\/jcb.117.6.1151","article-title":"Heat shock gene regulation by nascent polypeptides and denatured proteins: hsp70 as a potential autoregulatory factor","volume":"117","author":"Baler","year":"1992","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.mad.2026.112225_bib7","doi-asserted-by":"crossref","DOI":"10.3389\/fragi.2022.861761","article-title":"The CBP-1\/p300 lysine acetyltransferase regulates the heat shock response in C. elegans","volume":"3","author":"Barrett","year":"2022","journal-title":"Front. Aging"},{"key":"10.1016\/j.mad.2026.112225_bib8","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1038\/nrm4048","article-title":"Epigenetic regulation of ageing: linking environmental inputs to genomic stability","volume":"16","author":"Benayoun","year":"2015","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib9","doi-asserted-by":"crossref","first-page":"14914","DOI":"10.1073\/pnas.0902882106","article-title":"Collapse of proteostasis represents an early molecular event in Caenorhabditis elegans aging","volume":"106","author":"Ben-Zvi","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.mad.2026.112225_bib10","doi-asserted-by":"crossref","first-page":"8033","DOI":"10.1128\/MCB.19.12.8033","article-title":"Multiple components of the HSP90 chaperone complex function in regulation of heat shock factor 1 in vivo","volume":"19","author":"Bharadwaj","year":"1999","journal-title":"Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib11","doi-asserted-by":"crossref","first-page":"1471","DOI":"10.1002\/2211-5463.12484","article-title":"Induction of ubiquitin C ( UBC) gene transcription is mediated by HSF 1: role of proteotoxic and oxidative stress","volume":"8","author":"Bianchi","year":"2018","journal-title":"FEBS Open. Bio"},{"key":"10.1016\/j.mad.2026.112225_bib12","doi-asserted-by":"crossref","first-page":"2087","DOI":"10.4161\/cc.5.18.3288","article-title":"Aging and Immortality: Quasi-Programmed Senescence and Its Pharmacologic Inhibition","volume":"5","author":"Blagosklonny","year":"2006","journal-title":"Cell. Cycle"},{"key":"10.1016\/j.mad.2026.112225_bib13","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1186\/s12864-016-2837-5","article-title":"The genome-wide role of HSF-1 in the regulation of gene expression in Caenorhabditis elegans","volume":"17","author":"Brunquell","year":"2016","journal-title":"BMC Genom."},{"key":"10.1016\/j.mad.2026.112225_bib14","doi-asserted-by":"crossref","first-page":"113","DOI":"10.3390\/biom15010113","article-title":"Proteostasis decline and redox imbalance in age-related diseases: The therapeutic potential of NRF2","volume":"15","author":"Buttari","year":"2025","journal-title":"Biomolecules"},{"key":"10.1016\/j.mad.2026.112225_bib15","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1038\/nchembio.763","article-title":"Small-molecule proteostasis regulators for protein conformational diseases","volume":"8","author":"Calamini","year":"2011","journal-title":"Nat. Chem. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib16","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pbio.3003639","article-title":"Transcriptomic and chromatin accessibility profiling unveils new regulators of heat hormesis in Caenorhabditis elegans","volume":"24","author":"Chang","year":"2026","journal-title":"PLoS. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib17","doi-asserted-by":"crossref","first-page":"322","DOI":"10.1016\/j.cell.2011.12.019","article-title":"HSF-1 regulators DDL-1\/2 link insulin-like signaling to heat-shock responses and modulation of longevity","volume":"148","author":"Chiang","year":"2012","journal-title":"Cell"},{"key":"10.1016\/j.mad.2026.112225_bib18","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0039679","article-title":"mTOR is essential for the proteotoxic stress response, HSF1 activation and heat shock protein synthesis","volume":"7","author":"Chou","year":"2012","journal-title":"PLoS. One"},{"key":"10.1016\/j.mad.2026.112225_bib19","doi-asserted-by":"crossref","DOI":"10.1126\/science.1124646","article-title":"Opposing activities protect against age-onset proteotoxicity","volume":"313","author":"Cohen","year":"2006","journal-title":"Science"},{"key":"10.1016\/j.mad.2026.112225_bib20","doi-asserted-by":"crossref","first-page":"275","DOI":"10.15252\/embj.201489062","article-title":"Suppression of the HSF1-mediated proteotoxic stress response by the metabolic stress sensor AMPK","volume":"34","author":"Dai","year":"2015","journal-title":"EMBO J."},{"key":"10.1016\/j.mad.2026.112225_bib21","doi-asserted-by":"crossref","first-page":"1005","DOI":"10.1016\/j.cell.2007.07.020","article-title":"Heat shock factor 1 is a powerful multifaceted modifier of carcinogenesis","volume":"130","author":"Dai","year":"2007","journal-title":"Cell"},{"key":"10.1016\/j.mad.2026.112225_bib22","doi-asserted-by":"crossref","first-page":"139","DOI":"10.2174\/138920110790909704","article-title":"The heat shock connection of metabolic stress and dietary restriction","volume":"11","author":"Dancs\u00f3","year":"2010","journal-title":"Curr. Pharm. Biotechnol."},{"key":"10.1016\/j.mad.2026.112225_bib23","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1007038","article-title":"The homeodomain-interacting protein kinase HPK-1 preserves protein homeostasis and longevity through master regulatory control of the HSF-1 chaperone network and TORC1-restricted autophagy in Caenorhabditis elegans","volume":"13","author":"Das","year":"2017","journal-title":"PLoS. Genet."},{"key":"10.1016\/j.mad.2026.112225_bib24","doi-asserted-by":"crossref","first-page":"1606","DOI":"10.1111\/febs.13999","article-title":"Regulation of the mammalian heat shock factor 1","volume":"284","author":"Dayalan Naidu","year":"2017","journal-title":"FEBS J."},{"key":"10.1016\/j.mad.2026.112225_bib25","doi-asserted-by":"crossref","first-page":"2403","DOI":"10.1128\/MCB.00292-16","article-title":"Heat shock factor 1 is a substrate for p38 mitogen-activated protein kinases","volume":"36","author":"Dayalan Naidu","year":"2016","journal-title":"Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib26","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1016\/j.celrep.2015.07.026","article-title":"Heterotypic Signals from Neural HSF-1 Separate Thermotolerance from Longevity","volume":"12","author":"Douglas","year":"2015","journal-title":"Cell. Rep."},{"key":"10.1016\/j.mad.2026.112225_bib27","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2021.109623","article-title":"Insulin\/IGF-1 signaling and heat stress differentially regulate HSF1 activities in germline development","volume":"36","author":"Edwards","year":"2021","journal-title":"Cell. Rep."},{"key":"10.1016\/j.mad.2026.112225_bib28","doi-asserted-by":"crossref","DOI":"10.1016\/j.isci.2021.102951","article-title":"Structures of heat shock factor trimers bound to DNA","volume":"24","author":"Feng","year":"2021","journal-title":"iScience"},{"key":"10.1016\/j.mad.2026.112225_bib29","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1038\/s41556-019-0458-3","article-title":"HSF1 phase transition mediates stress adaptation and cell fate decisions","volume":"22","author":"Gaglia","year":"2020","journal-title":"Nat. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib30","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1038\/nrm.2017.73","article-title":"Regulation of heat shock transcription factors and their roles in physiology and disease","volume":"19","author":"Gomez-Pastor","year":"2018","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib31","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0243522","article-title":"Temporal requirements of SKN-1\/NRF as a regulator of lifespan and proteostasis in Caenorhabditis elegans","volume":"16","author":"Grushko","year":"2021","journal-title":"PLoS. ONE"},{"key":"10.1016\/j.mad.2026.112225_bib32","doi-asserted-by":"crossref","first-page":"1937","DOI":"10.1534\/genetics.104.028423","article-title":"The L-type cyclin CYL-1 and the heat-shock-factor HSF-1 are required for heat-shock-induced protein expression","volume":"168","author":"Hajdu-Cronin","year":"2004","journal-title":"Genetics"},{"key":"10.1016\/j.mad.2026.112225_bib33","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1038\/nature10317","article-title":"Molecular chaperones in protein folding and proteostasis","volume":"475","author":"Hartl","year":"2011","journal-title":"Nature"},{"key":"10.1016\/j.mad.2026.112225_bib34","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.11576","article-title":"Molecular mechanism of thermosensory function of human heat shock transcription factor Hsf1","volume":"5","author":"Hentze","year":"2016","journal-title":"eLife"},{"key":"10.1016\/j.mad.2026.112225_bib35","first-page":"2909","article-title":"Expression of heat shock protein 70 is altered by age and diet at the level of transcription","volume":"13","author":"Heydari","year":"1993","journal-title":"Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib36","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1002\/(SICI)1520-6408(1996)18:2<114::AID-DVG4>3.0.CO;2-C","article-title":"Effect of caloric restriction on the expression of heat shock protein 70 and the activation of heat shock transcription factor 1","volume":"18","author":"Heydari","year":"1996","journal-title":"Dev. Genet."},{"key":"10.1016\/j.mad.2026.112225_bib37","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1006\/excr.2000.4808","article-title":"Age-related alterations in the activation of heat shock transcription factor 1 in rat hepatocytes","volume":"256","author":"Heydari","year":"2000","journal-title":"Exp. Cell. Res."},{"key":"10.1016\/j.mad.2026.112225_bib38","doi-asserted-by":"crossref","first-page":"2522","DOI":"10.1091\/mbc.E18-06-0362","article-title":"Spatial regulation of the actin cytoskeleton by HSF-1 during aging","volume":"29","author":"Higuchi-Sanabria","year":"2018","journal-title":"MBoC"},{"key":"10.1016\/j.mad.2026.112225_bib39","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1038\/s41580-019-0101-y","article-title":"The proteostasis network and its decline in ageing","volume":"20","author":"Hipp","year":"2019","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib40","doi-asserted-by":"crossref","first-page":"40263","DOI":"10.1074\/jbc.M104714200","article-title":"Regulation of heat shock transcription factor 1 by stress-induced SUMO-1 modification","volume":"276","author":"Hong","year":"2001","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.mad.2026.112225_bib41","doi-asserted-by":"crossref","first-page":"1142","DOI":"10.1126\/science.1083701","article-title":"Regulation of aging and age-related disease by DAF-16 and heat-shock factor","volume":"300","author":"Hsu","year":"2003","journal-title":"Science"},{"key":"10.1016\/j.mad.2026.112225_bib42","article-title":"A default silencing mechanism restrains stress-induced genes in C. elegans","author":"Ilbay","year":"2025","journal-title":"bioRxiv"},{"key":"10.1016\/j.mad.2026.112225_bib43","doi-asserted-by":"crossref","first-page":"30459","DOI":"10.1074\/jbc.M114.591578","article-title":"Genomic heat shock element sequences drive cooperative human heat shock factor 1 DNA binding and selectivity","volume":"289","author":"Jaeger","year":"2014","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.mad.2026.112225_bib44","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1016\/j.celrep.2019.03.044","article-title":"Transcriptomics-based screening identifies pharmacological inhibition of Hsp90 as a means to defer aging","volume":"27","author":"Janssens","year":"2019","journal-title":"Cell. Rep."},{"key":"10.1016\/j.mad.2026.112225_bib45","doi-asserted-by":"crossref","first-page":"1406","DOI":"10.1038\/nm.4001","article-title":"Proteostasis and aging","volume":"21","author":"Kaushik","year":"2015","journal-title":"Nat. Med."},{"issue":"7288","key":"10.1016\/j.mad.2026.112225_bib46","doi-asserted-by":"crossref","first-page":"504","DOI":"10.1038\/nature08980","article-title":"The genetics of ageing","volume":"464","author":"Kenyon","year":"2010","journal-title":"Nature"},{"key":"10.1016\/j.mad.2026.112225_bib47","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1515\/bc.2010.058","article-title":"Chaperone-assisted degradation: multiple paths to destruction","volume":"391","author":"Kettern","year":"2010","journal-title":"Biol. Chem."},{"key":"10.1016\/j.mad.2026.112225_bib48","doi-asserted-by":"crossref","first-page":"617","DOI":"10.1111\/j.1474-9726.2012.00827.x","article-title":"Implications of time-series gene expression profiles of replicative senescence","volume":"11","author":"Kim","year":"2012","journal-title":"Aging Cell."},{"key":"10.1016\/j.mad.2026.112225_bib49","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.12821","article-title":"HSF-1 activates the ubiquitin proteasome system to promote non-apoptotic developmental cell death in C. elegans","volume":"5","author":"Kinet","year":"2016","journal-title":"eLife"},{"key":"10.1016\/j.mad.2026.112225_bib50","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1016\/j.tibs.2021.10.004","article-title":"Molecular mechanisms of heat shock factor 1 regulation","volume":"47","author":"Kmiecik","year":"2022","journal-title":"Trends Biochem. Sci."},{"key":"10.1016\/j.mad.2026.112225_bib51","doi-asserted-by":"crossref","DOI":"10.1111\/acel.14246","article-title":"Age-dependent heat shock hormesis to HSF-1 deficiency suggests a compensatory mechanism","volume":"23","author":"Kov\u00e1cs","year":"2024","journal-title":"Aging Cell."},{"key":"10.1016\/j.mad.2026.112225_bib52","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1007\/s42977-022-00138-z","article-title":"Functional diversification of heat shock factors","volume":"73","author":"Kov\u00e1cs","year":"2022","journal-title":"Biol. Futur."},{"key":"10.1016\/j.mad.2026.112225_bib53","doi-asserted-by":"crossref","first-page":"5815","DOI":"10.3390\/ijms20225815","article-title":"HSF1Base: A Comprehensive Database of HSF1 (Heat Shock Factor 1) Target Genes","volume":"20","author":"Kov\u00e1cs","year":"2019","journal-title":"IJMS"},{"key":"10.1016\/j.mad.2026.112225_bib54","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0077788","article-title":"Heat Shock Transcription Factor 1-Deficiency Attenuates Overloading-Associated Hypertrophy of Mouse Soleus Muscle","volume":"8","author":"Koya","year":"2013","journal-title":"PLoS. ONE"},{"key":"10.1016\/j.mad.2026.112225_bib55","doi-asserted-by":"crossref","first-page":"e31668","DOI":"10.7554\/eLife.31668","article-title":"Hsf1 and Hsp70 constitute a two-component feedback loop that regulates the yeast heat shock response","author":"Krakowiak","year":"2018","journal-title":"eLife"},{"key":"10.1016\/j.mad.2026.112225_bib56","doi-asserted-by":"crossref","first-page":"2177","DOI":"10.1152\/japplphysiol.01267.2001","article-title":"Invited Review: Heat shock proteins: modifying factors in physiological stress responses and acquired thermotolerance","volume":"92","author":"Kregel","year":"2002","journal-title":"J. Appl. Physiol."},{"key":"10.1016\/j.mad.2026.112225_bib57","doi-asserted-by":"crossref","first-page":"1950","DOI":"10.1016\/j.stem.2021.07.009","article-title":"Hsf1 promotes hematopoietic stem cell fitness and proteostasis in response to ex vivo culture stress and aging","volume":"28","author":"Kruta","year":"2021","journal-title":"Cell. Stem Cell."},{"key":"10.1016\/j.mad.2026.112225_bib58","doi-asserted-by":"crossref","DOI":"10.1038\/ncomms14337","article-title":"Hormetic heat stress and HSF-1 induce autophagy to improve survival and proteostasis in C. elegans","volume":"8","author":"Kumsta","year":"2017","journal-title":"Nat. Commun."},{"key":"10.1016\/j.mad.2026.112225_bib59","doi-asserted-by":"crossref","DOI":"10.1016\/j.ejmech.2021.113846","article-title":"The heat shock response and small molecule regulators","volume":"226","author":"Kurop","year":"2021","journal-title":"Eur. J. Med. Chem."},{"key":"10.1016\/j.mad.2026.112225_bib60","doi-asserted-by":"crossref","first-page":"14907","DOI":"10.3390\/ijms232314907","article-title":"The Thermal Stress Coping Network of the Nematode Caenorhabditis elegans","volume":"23","author":"Kyriakou","year":"2022","journal-title":"Int. J. Mol. Sci."},{"key":"10.1016\/j.mad.2026.112225_bib61","doi-asserted-by":"crossref","first-page":"1481","DOI":"10.1016\/j.celrep.2017.10.038","article-title":"Mitochondrial stress restores the heat shock response and prevents proteostasis collapse during aging","volume":"21","author":"Labbadia","year":"2017","journal-title":"Cell. Rep."},{"key":"10.1016\/j.mad.2026.112225_bib62","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1016\/j.molcel.2015.06.027","article-title":"Repression of the heat shock response is a programmed event at the onset of reproduction","volume":"59","author":"Labbadia","year":"2015","journal-title":"Mol. Cell."},{"key":"10.1016\/j.mad.2026.112225_bib63","doi-asserted-by":"crossref","DOI":"10.3389\/fragi.2022.861686","article-title":"HSF-1: Guardian of the proteome through integration of longevity signals to the proteostatic network","volume":"3","author":"Lazaro-Pena","year":"2022","journal-title":"Front. Aging"},{"key":"10.1016\/j.mad.2026.112225_bib64","doi-asserted-by":"crossref","first-page":"2454","DOI":"10.3390\/cells10092454","article-title":"Heat shock factor 1 prevents age-related hearing loss by decreasing endoplasmic reticulum stress","volume":"10","author":"Lee","year":"2021","journal-title":"Cells"},{"key":"10.1016\/j.mad.2026.112225_bib65","doi-asserted-by":"crossref","first-page":"2062","DOI":"10.1101\/gad.283317.116","article-title":"E2F coregulates an essential HSF developmental program that is distinct from the heat-shock response","volume":"30","author":"Li","year":"2016","journal-title":"Genes. Dev."},{"key":"10.1016\/j.mad.2026.112225_bib66","doi-asserted-by":"crossref","first-page":"2962","DOI":"10.1093\/nar\/gkp180","article-title":"Heat shock factor-1 modulates p53 activity in the transcriptional response to DNA damage","volume":"37","author":"Logan","year":"2009","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.mad.2026.112225_bib67","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1016\/j.cell.2022.11.001","article-title":"Hallmarks of aging: An expanding universe","volume":"186","author":"L\u00f3pez-Ot\u00edn","year":"2023","journal-title":"Cell"},{"key":"10.1016\/j.mad.2026.112225_bib68","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.molcel.2016.02.025","article-title":"Mammalian Heat Shock Response and Mechanisms Underlying Its Genome-wide Transcriptional Regulation","volume":"62","author":"Mahat","year":"2016","journal-title":"Mol. Cell."},{"key":"10.1016\/j.mad.2026.112225_bib69","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/0092-8674(95)90231-7","article-title":"Displacement of sequence-specific transcription factors from mitotic chromatin","volume":"83","author":"Martinez-Balbas","year":"1995","journal-title":"Cell"},{"key":"10.1016\/j.mad.2026.112225_bib70","doi-asserted-by":"crossref","DOI":"10.1016\/j.yexcr.2020.112246","article-title":"Hsf1 on a leash \u2014 controlling the heat shock response by chaperone titration","volume":"396","author":"Masser","year":"2020","journal-title":"Exp. Cell. Res."},{"key":"10.1016\/j.mad.2026.112225_bib71","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.47791","article-title":"Cytoplasmic protein misfolding titrates Hsp70 to activate nuclear Hsf1","volume":"8","author":"Masser","year":"2019","journal-title":"eLife"},{"key":"10.1016\/j.mad.2026.112225_bib72","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1016\/j.cell.2012.06.031","article-title":"HSF1 drives a transcriptional program distinct from heat shock to support highly malignant human cancers","volume":"150","author":"Mendillo","year":"2012","journal-title":"Cell"},{"key":"10.1016\/j.mad.2026.112225_bib73","doi-asserted-by":"crossref","first-page":"2765","DOI":"10.1242\/jcs.112.16.2765","article-title":"Human heat shock factor 1 is predominantly a nuclear protein before and after heat stress","volume":"112","author":"Mercier","year":"1999","journal-title":"J. Cell. Sci."},{"key":"10.1016\/j.mad.2026.112225_bib74","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pbio.3001605","article-title":"Transcellular chaperone signaling is an intercellular stress-response distinct from the HSF-1-mediated heat shock response","volume":"21","author":"Miles","year":"2023","journal-title":"PLoS. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib75","doi-asserted-by":"crossref","first-page":"1427","DOI":"10.1101\/gad.1657108","article-title":"Proteotoxic stress and inducible chaperone networks in neurodegenerative disease and aging","volume":"22","author":"Morimoto","year":"2008","journal-title":"Genes. Dev."},{"key":"10.1016\/j.mad.2026.112225_bib76","doi-asserted-by":"crossref","first-page":"S33","DOI":"10.1093\/gerona\/glu049","article-title":"Proteostasis and the aging proteome in health and disease","volume":"69","author":"Morimoto","year":"2014","journal-title":"J. Gerontol. A Biol. Sci. Med. Sci."},{"key":"10.1016\/j.mad.2026.112225_bib77","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1091\/mbc.e03-07-0532","article-title":"Regulation of Longevity in Caenorhabditis elegans by Heat Shock Factor and Molecular Chaperones","volume":"15","author":"Morley","year":"2004","journal-title":"MBoC"},{"key":"10.1016\/j.mad.2026.112225_bib78","doi-asserted-by":"crossref","DOI":"10.3389\/fragi.2022.899744","article-title":"Auxin-inducible degron system reveals temporal-spatial roles of HSF-1 and its transcriptional program in lifespan assurance","volume":"3","author":"Morphis","year":"2022","journal-title":"Front. Aging"},{"key":"10.1016\/j.mad.2026.112225_bib79","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1111\/acel.12024","article-title":"C. elegans HSF-1 is an essential nuclear protein that forms stress granule-like structures following heat shock","volume":"12","author":"Morton","year":"2013","journal-title":"Aging Cell."},{"issue":"21","key":"10.1016\/j.mad.2026.112225_bib80","doi-asserted-by":"crossref","first-page":"4892","DOI":"10.1038\/s44318-024-00234-x","article-title":"Non-cell-autonomous regulation of germline proteostasis by insulin\/IGF-1 signaling-induced dietary peptide uptake via PEPT-1","volume":"43","author":"Muhammad","year":"2024","journal-title":"EMBO J."},{"key":"10.1016\/j.mad.2026.112225_bib81","doi-asserted-by":"crossref","first-page":"1257","DOI":"10.1016\/S0531-5565(02)00134-1","article-title":"Chaperone function and chaperone overload in the aged. A preliminary analysis","volume":"37","author":"Nardai","year":"2002","journal-title":"Exp. Gerontol."},{"key":"10.1016\/j.mad.2026.112225_bib82","doi-asserted-by":"crossref","first-page":"703","DOI":"10.1016\/S0092-8674(03)00401-X","article-title":"Rb-mediated heterochromatin formation and silencing of E2F target genes during cellular senescence","volume":"113","author":"Narita","year":"2003","journal-title":"Cell"},{"key":"10.1016\/j.mad.2026.112225_bib83","article-title":"Modulation of heat shock transcription factor 1 as a therapeutic target for small molecule intervention in neurodegenerative disease","volume":"9","author":"Neef","year":"2011","journal-title":"PLoS. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib84","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1002673","article-title":"A Role for SKN-1\/Nrf in Pathogen Resistance and Immunosenescence in Caenorhabditis elegans","volume":"8","author":"Papp","year":"2012","journal-title":"PLoS. Pathog."},{"key":"10.1016\/j.mad.2026.112225_bib85","doi-asserted-by":"crossref","first-page":"C404","DOI":"10.1152\/ajpcell.00051.2002","article-title":"Redox signaling of cardiac HSF1 DNA binding","volume":"283","author":"Paroo","year":"2002","journal-title":"Am. J. Physiol. -Cell. Physiol."},{"key":"10.1016\/j.mad.2026.112225_bib86","doi-asserted-by":"crossref","first-page":"811","DOI":"10.1126\/science.1156093","article-title":"Regulation of the cellular heat shock response in Caenorhabditis elegans by thermosensory neurons","volume":"320","author":"Prahlad","year":"2008","journal-title":"Science"},{"key":"10.1016\/j.mad.2026.112225_bib87","doi-asserted-by":"crossref","first-page":"14204","DOI":"10.1073\/pnas.1106557108","article-title":"Neuronal circuitry regulates the response of Caenorhabditis elegans to misfolded proteins","volume":"108","author":"Prahlad","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.mad.2026.112225_bib88","doi-asserted-by":"crossref","first-page":"1046","DOI":"10.3390\/cells9041046","article-title":"HSF1: Primary factor in molecular chaperone expression and a major contributor to cancer morbidity","volume":"9","author":"Prince","year":"2020","journal-title":"Cells"},{"key":"10.1016\/j.mad.2026.112225_bib89","doi-asserted-by":"crossref","first-page":"723","DOI":"10.1083\/jcb.201607091","article-title":"The transcriptional regulator of the chaperone response HSF1 controls hepatic bioenergetics and protein homeostasis","volume":"216","author":"Qiao","year":"2017","journal-title":"J. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib90","doi-asserted-by":"crossref","first-page":"975","DOI":"10.1016\/j.cell.2014.01.055","article-title":"Interplay of acetyltransferase EP300 and the proteasome system in regulating heat shock transcription factor 1","volume":"156","author":"Raychaudhuri","year":"2014","journal-title":"Cell"},{"key":"10.1016\/j.mad.2026.112225_bib91","doi-asserted-by":"crossref","first-page":"8207","DOI":"10.1111\/febs.16178","article-title":"Interplay between mammalian heat shock factors 1 and 2 in physiology and pathology","volume":"289","author":"Roos-Mattjus","year":"2022","journal-title":"FEBS J."},{"key":"10.1016\/j.mad.2026.112225_bib92","doi-asserted-by":"crossref","first-page":"31902","DOI":"10.1073\/pnas.2018138117","article-title":"Cellular proteostasis decline in human senescence","volume":"117","author":"Sabath","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.mad.2026.112225_bib93","doi-asserted-by":"crossref","DOI":"10.1126\/science.1238303","article-title":"Tight Coordination of Protein Translation and HSF1 Activation Supports the Anabolic Malignant State","volume":"341","author":"Santagata","year":"2013","journal-title":"Science"},{"key":"10.1016\/j.mad.2026.112225_bib94","first-page":"1392","article-title":"Activation of heat shock gene transcription by heat shock factor 1 involves oligomerization, acquisition of DNA-binding activity, and nuclear localization","volume":"13","author":"Sarge","year":"1993","journal-title":"Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib95","doi-asserted-by":"crossref","first-page":"8984","DOI":"10.1038\/s41598-022-12736-x","article-title":"Binding of the HSF-1 DNA-binding domain to multimeric C. elegans consensus HSEs is guided by cooperative interactions","volume":"12","author":"Schmauder","year":"2022","journal-title":"Sci. Rep."},{"key":"10.1016\/j.mad.2026.112225_bib96","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1038\/nrm.2017.20","article-title":"The HSP90 chaperone machinery","volume":"18","author":"Schopf","year":"2017","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib97","doi-asserted-by":"crossref","first-page":"1073","DOI":"10.1111\/acel.12140","article-title":"Heat shock factor 1 mediates the longevity conferred by inhibition of TOR and insulin\/IGF-1 signaling pathways in C. elegans","volume":"12","author":"Seo","year":"2013","journal-title":"Aging Cell."},{"key":"10.1016\/j.mad.2026.112225_bib98","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1016\/j.tma.2020.07.002","article-title":"The heat shock transcription factor HSF-1 protects Caenorhabditis elegans from peroxide stress","volume":"4","author":"Servello","year":"2020","journal-title":"Transl. Med. Aging"},{"key":"10.1016\/j.mad.2026.112225_bib99","doi-asserted-by":"crossref","first-page":"1787","DOI":"10.1101\/gad.223834.113","article-title":"Lamin B1 depletion in senescent cells triggers large-scale changes in gene expression","volume":"27","author":"Shah","year":"2013","journal-title":"Genes. Dev."},{"key":"10.1016\/j.mad.2026.112225_bib100","doi-asserted-by":"crossref","first-page":"9872","DOI":"10.3390\/ijms26209872","article-title":"Beyond antioxidants: The emerging role of Nrf2 activation in amyotrophic lateral sclerosis (ALS)","volume":"26","author":"Sharbafshaaer","year":"2025","journal-title":"Int. J. Mol. Sci."},{"issue":"5","key":"10.1016\/j.mad.2026.112225_bib101","doi-asserted-by":"crossref","first-page":"654","DOI":"10.1101\/gad.12.5.654","article-title":"Molecularchaperones as HSF1-specific transcriptional\u00a0repressors","volume":"12","author":"Shi","year":"1998","journal-title":"Genes & Development"},{"key":"10.1016\/j.mad.2026.112225_bib102","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1007\/s12192-020-01074-4","article-title":"Age-dependent expression profiles of two adaptogenic systems and thermotolerance in Drosophila melanogaster","volume":"25","author":"Shilova","year":"2020","journal-title":"Cell. Stress. Chaperon-.-."},{"key":"10.1016\/j.mad.2026.112225_bib103","doi-asserted-by":"crossref","DOI":"10.1038\/s41598-018-30592-6","article-title":"DAF-21\/Hsp90 is required for C. elegans longevity by ensuring DAF-16\/FOXO isoform A function","volume":"8","author":"Somogyv\u00e1ri","year":"2018","journal-title":"Sci. Rep."},{"key":"10.1016\/j.mad.2026.112225_bib104","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1046\/j.1365-2435.2002.00639.x","article-title":"Decreased heat-shock resistance and down-regulation of Hsp70 expression with increasing age in adult Drosophila melanogaster","volume":"16","author":"S\u00f8rensen","year":"2002","journal-title":"Funct. Ecol."},{"key":"10.1016\/j.mad.2026.112225_bib105","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1016\/S0197-0186(02)00043-8","article-title":"Chaperones and aging: role in neurodegeneration and in other civilizational diseases","volume":"41","author":"S\u00f6ti","year":"2002","journal-title":"Neurochem. Int."},{"key":"10.1016\/j.mad.2026.112225_bib106","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1111\/j.1474-9726.2008.00385.x","article-title":"Dietary restriction suppresses proteotoxicity and enhances longevity by an HSF-1-dependent mechanism","volume":"7","author":"Steinkraus","year":"2008","journal-title":"Aging Cell."},{"key":"10.1016\/j.mad.2026.112225_bib107","doi-asserted-by":"crossref","first-page":"1679","DOI":"10.1534\/g3.119.400044","article-title":"HSB-1 inhibition and HSF-1 Overexpression trigger overlapping transcriptional changes to promote longevity in Caenorhabditis elegans","volume":"9","author":"Sural","year":"2019","journal-title":"G3 (Bethesda"},{"key":"10.1016\/j.mad.2026.112225_bib108","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1038\/nrm2918","article-title":"HSP90 at the hub of protein homeostasis: emerging mechanistic insights","volume":"11","author":"Taipale","year":"2010","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib109","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1016\/j.cub.2014.11.040","article-title":"Neuronal serotonin release triggers the heat shock response in C. elegans in the absence of temperature increase","volume":"25","author":"Tatum","year":"2015","journal-title":"Curr. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib110","doi-asserted-by":"crossref","first-page":"a004440","DOI":"10.1101\/cshperspect.a004440","article-title":"Aging as an event of proteostasis collapse","volume":"3","author":"Taylor","year":"2011","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib111","doi-asserted-by":"crossref","first-page":"E3388","DOI":"10.1073\/pnas.1305275110","article-title":"Transcriptional response to stress in the dynamic chromatin environment of cycling and mitotic cells","volume":"110","author":"Vihervaara","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.mad.2026.112225_bib112","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1007\/978-0-387-39975-1_9","article-title":"Chaperone regulation of the heat shock protein response","volume":"594","author":"Voellmy","year":"2007","journal-title":"Adv. Exp. Med. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib113","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1111\/j.1474-9726.2012.00811.x","article-title":"Temporal requirements of heat shock factor-1 for longevity assurance","volume":"11","author":"Volovik","year":"2012","journal-title":"Aging Cell."},{"key":"10.1016\/j.mad.2026.112225_bib114","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1111\/j.1600-0854.2005.00266.x","article-title":"Constitutive nuclear import and stress-regulated nucleocytoplasmic shuttling of mammalian heat-shock factor 1","volume":"6","author":"Vujanac","year":"2005","journal-title":"Traffic"},{"key":"10.1016\/j.mad.2026.112225_bib115","article-title":"Reduced proteasome degradation of HSF-1 shifts protein stress management with age in Caenorhabditis elegans","volume":"25","author":"Wang","year":"2026","journal-title":"Aging Cell."},{"key":"10.1016\/j.mad.2026.112225_bib116","doi-asserted-by":"crossref","first-page":"13804","DOI":"10.3390\/ijms241813804","article-title":"Roles of stress response in autophagy processes and aging-related diseases","volume":"24","author":"Watanabe","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"10.1016\/j.mad.2026.112225_bib117","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1126\/science.1165946","article-title":"Stress-inducible regulation of heat shock factor 1 by the deacetylase SIRT1","volume":"323","author":"Westerheide","year":"2009","journal-title":"Science"},{"key":"10.1016\/j.mad.2026.112225_bib118","doi-asserted-by":"crossref","first-page":"758","DOI":"10.1016\/j.devcel.2020.06.038","article-title":"A mitochondrial stress-specific form of HSF1 protects against age-related proteostasis collapse","volume":"54","author":"Williams","year":"2020","journal-title":"Dev. Cell."},{"key":"10.1016\/j.mad.2026.112225_bib119","doi-asserted-by":"crossref","DOI":"10.1038\/cddis.2016.22","article-title":"The transcriptional coactivator PGC1\u03b1 protects against hyperthermic stress via cooperation with the heat shock factor HSF1","volume":"7","author":"Xu","year":"2016","journal-title":"Cell. Death Dis."},{"issue":"5","key":"10.1016\/j.mad.2026.112225_bib120","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1089\/omi.2011.0118","article-title":"clusterProfiler: an R Package for Comparing Biological Themes Among Gene Clusters","volume":"16","author":"Yu","year":"2012","journal-title":"OMICS: J. Integr. Biol."},{"key":"10.1016\/j.mad.2026.112225_bib121","doi-asserted-by":"crossref","first-page":"3351","DOI":"10.7150\/ijbs.110447","article-title":"HSF1 activation mechanisms, disease roles, and small molecule therapeutics","volume":"21","author":"Zhang","year":"2025","journal-title":"Int. J. Biol. Sci."},{"key":"10.1016\/j.mad.2026.112225_bib122","article-title":"Skeletal muscle HSF1 alleviates age-associated sarcopenia and mitochondrial function decline via SIRT3-PGC1\u03b1axis","volume":"13","author":"Zhang","year":"2026","journal-title":"Adv. Sci."},{"key":"10.1016\/j.mad.2026.112225_bib123","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1008122","article-title":"Histone acetylation promotes long-lasting defense responses and longevity following early life heat stress","volume":"15","author":"Zhou","year":"2019","journal-title":"PLoS. Genet."},{"key":"10.1016\/j.mad.2026.112225_bib124","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1016\/S0092-8674(00)81588-3","article-title":"Repression of Heat Shock Transcription Factor HSF1 Activation by HSP90 (HSP90 Complex) that Forms a Stress-Sensitive Complex with HSF1","volume":"94","author":"Zou","year":"1998","journal-title":"Cell"}],"container-title":["Mechanisms of Ageing and Development"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0047637426000771?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0047637426000771?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,11,9]],"date-time":"2026-11-09T21:04:27Z","timestamp":1794258267000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0047637426000771"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10]]},"references-count":124,"alternative-id":["S0047637426000771"],"URL":"https:\/\/doi.org\/10.1016\/j.mad.2026.112225","relation":{},"ISSN":["0047-6374"],"issn-type":[{"value":"0047-6374","type":"print"}],"subject":[],"published":{"date-parts":[[2026,10]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"HSF1 in aging: The output selection model","name":"articletitle","label":"Article Title"},{"value":"Mechanisms of Ageing and Development","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.mad.2026.112225","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Author(s). Published by Elsevier B.V.","name":"copyright","label":"Copyright"}],"article-number":"112225"}}