{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T21:00:44Z","timestamp":1775077244772,"version":"3.50.1"},"reference-count":84,"publisher":"Oxford University Press (OUP)","issue":"4","license":[{"start":{"date-parts":[[2026,3,3]],"date-time":"2026-03-03T00:00:00Z","timestamp":1772496000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN-2017-06738"],"award-info":[{"award-number":["RGPIN-2017-06738"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100018624","name":"USRA","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100018624","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,4,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Caenorhabditis elegans insulin\/insulin-like growth factor 1 signaling, IIS, affects diverse physiological processes through the DAF-16\/FOXO transcription factor. Despite its ubiquitous presence in somatic cells, DAF-16's effects exhibit prevalent tissue specificity as well as tissue crosstalk. This implies that tissue-specific DAF-16 transcriptional programs contribute to functional diversity of IIS. To further investigate this possibility, we sought muscle-cell-specific DAF-16 transcriptional targets. Using fluorescence-activated cell sorting to enrich for body wall muscle cells from young hermaphroditic adults, we compared the muscle cell mRNA transcriptomes under conditions of high and low IIS activity, with and without DAF-16. We further analyzed DAF-16a's binding sites in muscle and intestine cells by chromatin-immunoprecipitation sequencing. Combined output of these analyses is 12 candidate DAF-16 targets enriched for muscle cells. Transcriptional and translational reporters for three out of the four top candidates\u2014a secreted protein C54F6.5, a calcium-binding protein CEX-1\/calexcitin, and a fatty acid metabolic enzyme MLCD-1\/MCD\u2014showed DAF-16-dependent activation specifically in body wall muscle cells. Notably, reporters for C54F6.5 and cex-1 exhibit DAF-16-independent, constitutive expression in nonmuscle cells, explaining their low rank or absence from the DAF-16 target lists generated by whole-animal microarray or mRNA-sequencing analyses. These results highlight the need to examine FOXO targets in a cell-type-specific manner.<\/jats:p>","DOI":"10.1093\/g3journal\/jkag030","type":"journal-article","created":{"date-parts":[[2026,3,3]],"date-time":"2026-03-03T08:29:41Z","timestamp":1772526581000},"source":"Crossref","is-referenced-by-count":0,"title":["Three muscle-specific DAF-16\/FOXO transcriptional targets activated by reduced insulin\/IGF-1 signaling"],"prefix":"10.1093","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9158-9695","authenticated-orcid":false,"given":"Shifei","family":"Wu","sequence":"first","affiliation":[{"name":"Department of Molecular Genetics, University of Toronto , Toronto, ON M5S 3K3 ,","place":["Canada"]},{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jinghua","family":"Gao","sequence":"additional","affiliation":[{"name":"Department of Molecular Genetics, University of Toronto , Toronto, ON M5S 3K3 ,","place":["Canada"]},{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yan","family":"Li","sequence":"additional","affiliation":[{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8955-2423","authenticated-orcid":false,"given":"Charline","family":"Roy","sequence":"additional","affiliation":[{"name":"Universite\u0301 Claude Bernard Lyon 1, CNRS UMR 5284, INSERM U1314, MeLiS, 69008 Lyon ,","place":["France"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ying","family":"Wang","sequence":"additional","affiliation":[{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3336-245X","authenticated-orcid":false,"given":"Ben","family":"Mulcahy","sequence":"additional","affiliation":[{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"William","family":"Li","sequence":"additional","affiliation":[{"name":"Department of Molecular Genetics, University of Toronto , Toronto, ON M5S 3K3 ,","place":["Canada"]},{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sruthy","family":"Ravivarma","sequence":"additional","affiliation":[{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]},{"name":"Department of Cell and Systems Biology, University of Toronto , Toronto, ON M5S 3G5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2197-7801","authenticated-orcid":false,"given":"John","family":"Calarco","sequence":"additional","affiliation":[{"name":"Department of Cell and Systems Biology, University of Toronto , Toronto, ON M5S 3G5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8488-7638","authenticated-orcid":false,"given":"Wesley","family":"Hung","sequence":"additional","affiliation":[{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0086-9622","authenticated-orcid":false,"given":"Mei","family":"Zhen","sequence":"additional","affiliation":[{"name":"Department of Molecular Genetics, University of Toronto , Toronto, ON M5S 3K3 ,","place":["Canada"]},{"name":"Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital , Toronto, ON M5G 1X5 ,","place":["Canada"]},{"name":"Department of Cell and Systems Biology, University of Toronto , Toronto, ON M5S 3G5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2026,3,3]]},"reference":[{"key":"2026040114291463700_jkag030-B1","doi-asserted-by":"publisher","first-page":"e3001204","DOI":"10.1371\/journal.pbio.3001204","article-title":"DAF-16\/FoxO and DAF-12\/VDR control cellular plasticity both cell-autonomously and via interorgan signaling","volume":"19","author":"Aghayeva","year":"2021","journal-title":"PLoS Biol"},{"key":"2026040114291463700_jkag030-B2","doi-asserted-by":"publisher","first-page":"529","DOI":"10.1016\/s0166-2236(98)01277-6","article-title":"Time domains of neuronal Ca2+ signaling and associative memory: steps through a calexcitin, ryanodine receptor, K+ channel cascade","volume":"21","author":"Alkon","year":"1998","journal-title":"Trends Neurosci"},{"key":"2026040114291463700_jkag030-B3","doi-asserted-by":"publisher","first-page":"e1005788","DOI":"10.1371\/journal.pgen.1005788","article-title":"DAF-16 and TCER-1 facilitate adaptation to germline loss by restoring lipid homeostasis and repressing reproductive physiology in C. elegans","volume":"12","author":"Amrit","year":"2016","journal-title":"PLoS Genet"},{"key":"2026040114291463700_jkag030-B4","doi-asserted-by":"publisher","first-page":"D344","DOI":"10.1093\/nar\/gkaa977","article-title":"The InterPro protein families and domains database: 20 years on","volume":"49","author":"Blum","year":"2021","journal-title":"Nucleic Acids Res"},{"key":"2026040114291463700_jkag030-B5","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1093\/genetics\/77.1.71","article-title":"The genetics of Caenorhabditis elegans","volume":"77","author":"Brenner","year":"1974","journal-title":"Genetics"},{"key":"2026040114291463700_jkag030-B6","doi-asserted-by":"publisher","first-page":"561","DOI":"10.1016\/j.neuron.2013.11.019","article-title":"Insulin\/IGF1 signaling inhibits age-dependent axon regeneration","volume":"81","author":"Byrne","year":"2014","journal-title":"Neuron"},{"key":"2026040114291463700_jkag030-B8","doi-asserted-by":"publisher","first-page":"110330","DOI":"10.1016\/j.celrep.2022.110330","article-title":"Escape steering by cholecystokinin peptidergic signaling","volume":"38","author":"Chen","year":"2022","journal-title":"Cell Rep"},{"key":"2026040114291463700_jkag030-B7","doi-asserted-by":"publisher","first-page":"572","DOI":"10.1016\/j.neuron.2012.11.025","article-title":"Two insulin-like peptides antagonistically regulate aversive olfactory learning in C. elegans","volume":"77","author":"Chen","year":"2013","journal-title":"Neuron"},{"key":"2026040114291463700_jkag030-B9","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1186\/1749-8104-5-7","article-title":"Insulin receptor signaling in the development of neuronal structure and function","volume":"5","author":"Chiu","year":"2010","journal-title":"Neural Dev"},{"key":"2026040114291463700_jkag030-B10","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1016\/j.cmet.2009.02.003","article-title":"Coordinated regulation of foraging and metabolism in C. elegans by RFamide neuropeptide signaling","volume":"9","author":"Cohen","year":"2009","journal-title":"Cell Metab"},{"key":"2026040114291463700_jkag030-B11","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1038\/s41586-019-1524-5","article-title":"The flight response impairs cytoprotective mechanisms by activating the insulin pathway","volume":"573","author":"De Rosa","year":"2019","journal-title":"Nature"},{"key":"2026040114291463700_jkag030-B12","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1016\/j.ydbio.2008.03.019","article-title":"Insulin-like signaling negatively regulates muscle arm extension through DAF-12 in Caenorhabditis elegans","volume":"318","author":"Dixon","year":"2008","journal-title":"Dev Biol"},{"key":"2026040114291463700_jkag030-B13","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1093\/genetics\/159.1.133","article-title":"Genetic analysis of endocytosis in Caenorhabditis elegans: coelomocyte uptake defective mutants","volume":"159","author":"Fares","year":"2001","journal-title":"Genetics"},{"key":"2026040114291463700_jkag030-B14","doi-asserted-by":"publisher","first-page":"e1004225","DOI":"10.1371\/journal.pgen.1004225","article-title":"An insulin-to-insulin regulatory network orchestrates phenotypic specificity in development and physiology","volume":"10","author":"Fernandes de Abreu","year":"2014","journal-title":"PLoS Genet"},{"key":"2026040114291463700_jkag030-B15","doi-asserted-by":"publisher","first-page":"2149","DOI":"10.1101\/gad.1701508","article-title":"C. elegans dauer formation and the molecular basis of plasticity","volume":"22","author":"Fielenbach","year":"2008","journal-title":"Genes Dev"},{"key":"2026040114291463700_jkag030-B16","doi-asserted-by":"publisher","first-page":"22529","DOI":"10.1074\/jbc.M010508200","article-title":"Calexcitin B is a new member of the sarcoplasmic calcium-binding protein family","volume":"276","author":"Gombos","year":"2001","journal-title":"J Biol Chem"},{"key":"2026040114291463700_jkag030-B17","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1016\/j.cell.2015.02.018","article-title":"Feedback from network states generates variability in a probabilistic olfactory circuit","volume":"161","author":"Gordus","year":"2015","journal-title":"Cell"},{"key":"2026040114291463700_jkag030-B18","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1093\/genetics\/137.1.107","article-title":"daf-2, daf-16 and daf-23: genetically interacting genes controlling Dauer formation in Caenorhabditis elegans","volume":"137","author":"Gottlieb","year":"1994","journal-title":"Genetics"},{"key":"2026040114291463700_jkag030-B19","doi-asserted-by":"publisher","first-page":"8919","DOI":"10.1038\/ncomms9919","article-title":"C. elegans maximum velocity correlates with healthspan and is maintained in worms with an insulin receptor mutation","volume":"6","author":"Hahm","year":"2015","journal-title":"Nat Commun"},{"key":"2026040114291463700_jkag030-B20","doi-asserted-by":"publisher","first-page":"1975","DOI":"10.1016\/s0960-9822(01)00594-2","article-title":"daf-16 integrates developmental and environmental inputs to mediate aging in the nematode Caenorhabditis elegans","volume":"11","author":"Henderson","year":"2001","journal-title":"Curr Biol"},{"key":"2026040114291463700_jkag030-B21","doi-asserted-by":"publisher","first-page":"e30057","DOI":"10.7554\/eLife.30057","article-title":"daf-16\/FoxO promotes gluconeogenesis and trehalose synthesis during starvation to support survival","volume":"6","author":"Hibshman","year":"2017","journal-title":"eLife"},{"key":"2026040114291463700_jkag030-B22","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1186\/1471-2199-9-9","article-title":"Selection and validation of a set of reliable reference genes for quantitative sod gene expression analysis in C. elegans","volume":"9","author":"Hoogewijs","year":"2008","journal-title":"BMC Mol Biol"},{"key":"2026040114291463700_jkag030-B23","doi-asserted-by":"publisher","first-page":"1745","DOI":"10.1038\/emboj.2013.91","article-title":"Attenuation of insulin signalling contributes to FSN-1-mediated regulation of synapse development","volume":"32","author":"Hung","year":"2013","journal-title":"EMBO J"},{"key":"2026040114291463700_jkag030-B24","doi-asserted-by":"publisher","first-page":"1767","DOI":"10.1242\/dev.103846","article-title":"A Caenorhabditis elegans developmental decision requires insulin signaling-mediated neuron-intestine communication","volume":"141","author":"Hung","year":"2014","journal-title":"Development"},{"key":"2026040114291463700_jkag030-B25","doi-asserted-by":"publisher","first-page":"e68848","DOI":"10.7554\/eLife.68848","article-title":"Corollary discharge promotes a sustained motor state in a neural circuit for navigation","volume":"10","author":"Ji","year":"2021","journal-title":"eLife"},{"key":"2026040114291463700_jkag030-B26","doi-asserted-by":"publisher","first-page":"632","DOI":"10.1016\/j.cell.2016.01.007","article-title":"Distinct circuits for the formation and retrieval of an imprinted olfactory memory","volume":"164","author":"Jin","year":"2016","journal-title":"Cell"},{"key":"2026040114291463700_jkag030-B27","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1038\/nature16483","article-title":"The C. elegans adult neuronal IIS\/FOXO transcriptome reveals adult phenotype regulators","volume":"529","author":"Kaletsky","year":"2016","journal-title":"Nature"},{"key":"2026040114291463700_jkag030-B28","doi-asserted-by":"publisher","first-page":"415","DOI":"10.1242\/dmm.001040","article-title":"The role of insulin\/IGF-like signaling in C. elegans longevity and aging","volume":"3","author":"Kaletsky","year":"2010","journal-title":"Dis Model Mech"},{"key":"2026040114291463700_jkag030-B29","doi-asserted-by":"publisher","first-page":"e1005731","DOI":"10.1371\/journal.pgen.1005731","article-title":"dbl-1\/TGF-\u03b2 and daf-12\/NHR signaling mediate cell-nonautonomous effects of daf-16\/FOXO on starvation-induced developmental arrest","volume":"11","author":"Kaplan","year":"2015","journal-title":"PLOS Genet"},{"key":"2026040114291463700_jkag030-B30","doi-asserted-by":"publisher","first-page":"504","DOI":"10.1038\/nature08980","article-title":"The genetics of ageing","volume":"464","author":"Kenyon","year":"2010","journal-title":"Nature"},{"key":"2026040114291463700_jkag030-B31","doi-asserted-by":"publisher","first-page":"461","DOI":"10.1038\/366461a0","article-title":"A C. elegans mutant that lives twice as long as wild type","volume":"366","author":"Kenyon","year":"1993","journal-title":"Nature"},{"key":"2026040114291463700_jkag030-B32","doi-asserted-by":"publisher","first-page":"942","DOI":"10.1126\/science.277.5328.942","article-title":"daf-2, an insulin receptor-like gene that regulates longevity and diapause in Caenorhabditis elegans","volume":"277","author":"Kimura","year":"1997","journal-title":"Science"},{"key":"2026040114291463700_jkag030-B33","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1016\/j.mce.2008.10.048","article-title":"The endocrine regulation of aging in Caenorhabditis elegans","volume":"299","author":"Kleemann","year":"2009","journal-title":"Mol Cell Endocrinol"},{"key":"2026040114291463700_jkag030-B34","doi-asserted-by":"publisher","first-page":"911","DOI":"10.1038\/nn.2847","article-title":"High-fat feeding promotes obesity via insulin receptor\/PI3K-dependent inhibition of SF-1 VMH neurons","volume":"14","author":"Kl\u00f6ckener","year":"2011","journal-title":"Nat Neurosci"},{"key":"2026040114291463700_jkag030-B35","doi-asserted-by":"publisher","first-page":"2955","DOI":"10.1101\/gad.1479906","article-title":"Insulin-like signaling and the neural circuit for integrative behavior in C. elegans","volume":"20","author":"Kodama","year":"2006","journal-title":"Genes Dev"},{"key":"2026040114291463700_jkag030-B36","doi-asserted-by":"publisher","first-page":"937","DOI":"10.1534\/genetics.117.300657","article-title":"The ModERN resource: genome-wide binding profiles for hundreds of Drosophila and Caenorhabditis elegans transcription factors","volume":"208","author":"Kudron","year":"2018","journal-title":"Genetics"},{"key":"2026040114291463700_jkag030-B37","doi-asserted-by":"publisher","first-page":"1950","DOI":"10.1016\/s0960-9822(01)00595-4","article-title":"Regulation of C. elegans DAF-16 and its human ortholog FKHRL1 by the daf-2 insulin-like signaling pathway","volume":"11","author":"Lee","year":"2001","journal-title":"Curr Biol"},{"key":"2026040114291463700_jkag030-B38","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1016\/j.cmet.2009.10.003","article-title":"Glucose shortens the life span of C. elegans by downregulating DAF-16\/FOXO activity and aquaporin gene expression","volume":"10","author":"Lee","year":"2009","journal-title":"Cell Metab"},{"key":"2026040114291463700_jkag030-B39","doi-asserted-by":"publisher","first-page":"e12896","DOI":"10.1111\/acel.12896","article-title":"DAF-16 stabilizes the aging transcriptome and is activated in mid-aged Caenorhabditis elegans to cope with internal stress","volume":"18","author":"Li","year":"2019","journal-title":"Aging Cell"},{"key":"2026040114291463700_jkag030-B40","doi-asserted-by":"publisher","first-page":"844","DOI":"10.1101\/gad.1066503","article-title":"daf-28 encodes a C. elegans insulin superfamily member that is regulated by environmental cues and acts in the DAF-2 signaling pathway","volume":"17","author":"Li","year":"2003","journal-title":"Genes Dev"},{"key":"2026040114291463700_jkag030-B41","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1895\/wormbook.1.142.1","article-title":"Neuropeptides","author":"Li","year":"2008","journal-title":"WormBook"},{"key":"2026040114291463700_jkag030-B42","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1016\/S0092-8674(03)00889-4","article-title":"Tissue-specific activities of C. elegans DAF-16 in the regulation of lifespan","volume":"115","author":"Libina","year":"2003","journal-title":"Cell"},{"key":"2026040114291463700_jkag030-B43","doi-asserted-by":"publisher","first-page":"8001","DOI":"10.1523\/JNEUROSCI.4636-09.2010","article-title":"Insulin signaling plays a dual role in Caenorhabditis elegans memory acquisition and memory retrieval","volume":"30","author":"Lin","year":"2010","journal-title":"J Neurosci Off J Soc Neurosci"},{"key":"2026040114291463700_jkag030-B44","doi-asserted-by":"publisher","first-page":"392","DOI":"10.1016\/j.cmet.2013.08.007","article-title":"Functional aging in the nervous system contributes to age-dependent motor activity decline in C. elegans","volume":"18","author":"Liu","year":"2013","journal-title":"Cell Metab"},{"key":"2026040114291463700_jkag030-B45","doi-asserted-by":"publisher","first-page":"3878","DOI":"10.1038\/onc.2013.347","article-title":"Insulin activates the insulin receptor to downregulate the PTEN tumour suppressor","volume":"33","author":"Liu","year":"2014","journal-title":"Oncogene"},{"key":"2026040114291463700_jkag030-B46","doi-asserted-by":"publisher","first-page":"649","DOI":"10.1016\/s0896-6273(00)81067-3","article-title":"Regulation of AMPA receptor-mediated synaptic transmission by clathrin-dependent receptor internalization","volume":"25","author":"Man","year":"2000","journal-title":"Neuron"},{"key":"2026040114291463700_jkag030-B47","doi-asserted-by":"publisher","first-page":"260","DOI":"10.1016\/j.cmet.2010.08.004","article-title":"Insulin-like signaling determines survival during stress via post transcriptional mechanisms in C. elegans","volume":"12","author":"McColl","year":"2010","journal-title":"Cell Metab"},{"key":"2026040114291463700_jkag030-B48","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1046\/j.1474-9728.2003.00043.x","article-title":"Transcriptional outputs of the Caenorhabditis elegans forkhead protein DAF-16","volume":"2","author":"McElwee","year":"2003","journal-title":"Aging Cell"},{"key":"2026040114291463700_jkag030-B49","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1016\/S0091-679X(08)61399-0","article-title":"DNA transformation","volume":"48","author":"Mello","year":"1995","journal-title":"Methods Cell Biol"},{"key":"2026040114291463700_jkag030-B50","doi-asserted-by":"publisher","first-page":"1568","DOI":"10.1111\/j.1471-4159.2005.03155.x","article-title":"A biochemical and functional characterization of diet-induced brain insulin resistance","volume":"93","author":"Mielke","year":"2005","journal-title":"J Neurochem"},{"key":"2026040114291463700_jkag030-B51","doi-asserted-by":"publisher","first-page":"1985","DOI":"10.1534\/genetics.117.202747","article-title":"Heat-induced calcium leakage causes mitochondrial damage in Caenorhabditis elegans body-wall muscles","volume":"206","author":"Momma","year":"2017","journal-title":"Genetics"},{"key":"2026040114291463700_jkag030-B52","doi-asserted-by":"publisher","first-page":"536","DOI":"10.1038\/382536a0","article-title":"A phosphatidylinositol-3-OH kinase family member regulating longevity and diapause in Caenorhabditis elegans","volume":"382","author":"Morris","year":"1996","journal-title":"Nature"},{"key":"2026040114291463700_jkag030-B53","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1038\/nature01789","article-title":"Genes that act downstream of DAF-16 to influence the lifespan of Caenorhabditis elegans","volume":"424","author":"Murphy","year":"2003","journal-title":"Nature"},{"key":"2026040114291463700_jkag030-B54","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1895\/wormbook.1.164.1","article-title":"Insulin\/insulin-like growth factor signaling in C. elegans","author":"Murphy","year":"2013","journal-title":"WormBook"},{"key":"2026040114291463700_jkag030-B55","doi-asserted-by":"publisher","first-page":"13808","DOI":"10.1073\/pnas.93.24.13808","article-title":"Calexcitin: a signaling protein that binds calcium and GTP, inhibits potassium channels, and enhances membrane excitability","volume":"93","author":"Nelson","year":"1996","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026040114291463700_jkag030-B56","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1042\/0264-6021:3410423","article-title":"Calexcitin interaction with neuronal ryanodine receptors","volume":"341","author":"Nelson","year":"1999","journal-title":"Biochem J"},{"key":"2026040114291463700_jkag030-B57","doi-asserted-by":"publisher","first-page":"52","DOI":"10.3389\/fnins.2016.00052","article-title":"IGF-I: a key growth factor that regulates neurogenesis and synaptogenesis from embryonic to adult stages of the brain","volume":"10","author":"Nieto-Est\u00e9vez","year":"2016","journal-title":"Front Neurosci"},{"key":"2026040114291463700_jkag030-B58","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1152\/jn.01029.2010","article-title":"Neuronal plasticity regulated by the insulin-like signaling pathway underlies salt chemotaxis learning in Caenorhabditis elegans","volume":"106","author":"Oda","year":"2011","journal-title":"J Neurophysiol"},{"key":"2026040114291463700_jkag030-B59","doi-asserted-by":"publisher","first-page":"994","DOI":"10.1038\/40194","article-title":"The Fork head transcription factor DAF-16 transduces insulin-like metabolic and longevity signals in C. elegans","volume":"389","author":"Ogg","year":"1997","journal-title":"Nature"},{"key":"2026040114291463700_jkag030-B60","doi-asserted-by":"publisher","first-page":"19024","DOI":"10.1073\/pnas.1308866110","article-title":"Reduced IGF signaling prevents muscle cell death in a Caenorhabditis elegans model of muscular dystrophy","volume":"110","author":"Oh","year":"2013","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026040114291463700_jkag030-B61","doi-asserted-by":"publisher","first-page":"9274","DOI":"10.1073\/pnas.1011711108","article-title":"Genetic analysis of age-dependent defects of the Caenorhabditis elegans touch receptor neurons","volume":"108","author":"Pan","year":"2011","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026040114291463700_jkag030-B62","doi-asserted-by":"publisher","first-page":"1438","DOI":"10.1101\/gad.13.11.1438","article-title":"A PDK1 homolog is necessary and sufficient to transduce AGE-1 PI3 kinase signals that regulate diapause in Caenorhabditis elegans","volume":"13","author":"Paradis","year":"1999","journal-title":"Genes Dev"},{"key":"2026040114291463700_jkag030-B63","doi-asserted-by":"publisher","first-page":"2488","DOI":"10.1101\/gad.12.16.2488","article-title":"Caenorhabditis elegans Akt\/PKB transduces insulin receptor-like signals from AGE-1 PI3 kinase to the DAF-16 transcription factor","volume":"12","author":"Paradis","year":"1998","journal-title":"Genes Dev"},{"key":"2026040114291463700_jkag030-B64","doi-asserted-by":"publisher","first-page":"672","DOI":"10.1101\/gad.867301","article-title":"Regulation of DAF-2 receptor signaling by human insulin and ins-1, a member of the unusually large and diverse C. elegans insulin gene family","volume":"15","author":"Pierce","year":"2001","journal-title":"Genes Dev"},{"key":"2026040114291463700_jkag030-B65","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1016\/j.biosystems.2008.06.003","article-title":"Conservation of key members in the course of the evolution of the insulin signaling pathway","volume":"95","author":"Pi\u00f1ero Gonz\u00e1lez","year":"2009","journal-title":"Biosystems"},{"key":"2026040114291463700_jkag030-B66","doi-asserted-by":"publisher","first-page":"1886","DOI":"10.1172\/JCI27123","article-title":"Enhanced PIP3 signaling in POMC neurons causes KATP channel activation and leads to diet-sensitive obesity","volume":"116","author":"Plum","year":"2006","journal-title":"J Clin Invest"},{"key":"2026040114291463700_jkag030-B67","doi-asserted-by":"publisher","first-page":"2052","DOI":"10.1161\/ATVBAHA.111.241919","article-title":"Tissue-specific insulin signaling, metabolic syndrome, and cardiovascular disease","volume":"32","author":"Rask-Madsen","year":"2012","journal-title":"Arterioscler Thromb Vasc Biol"},{"key":"2026040114291463700_jkag030-B68","doi-asserted-by":"publisher","first-page":"105","DOI":"10.3390\/metabo11020105","article-title":"Elevated trehalose levels in C. elegans daf-2 mutants increase stress resistance, not lifespan","volume":"11","author":"Rasulova","year":"2021","journal-title":"Metabolites"},{"issue":"8","key":"2026040114291463700_jkag030-B69","doi-asserted-by":"publisher","first-page":"e13660","DOI":"10.1111\/acel.13660","article-title":"DAF-2\/insulin IGF-1 receptor regulates motility during aging by integrating opposite signaling from muscle and neuronal tissues","volume":"21","author":"Roy","year":"2022","journal-title":"Aging Cell"},{"key":"2026040114291463700_jkag030-B70","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat Methods"},{"key":"2026040114291463700_jkag030-B71","doi-asserted-by":"publisher","first-page":"399","DOI":"10.1038\/msb.2010.54","article-title":"DamID in C. elegans reveals longevity-associated targets of DAF-16\/FoxO","volume":"6","author":"Schuster","year":"2010","journal-title":"Mol Syst Biol"},{"key":"2026040114291463700_jkag030-B72","doi-asserted-by":"publisher","first-page":"e67723","DOI":"10.7554\/eLife.67723","article-title":"Behavioral control by depolarized and hyperpolarized states of an integrating neuron","volume":"10","author":"Sordillo","year":"2021","journal-title":"eLife"},{"key":"2026040114291463700_jkag030-B73","doi-asserted-by":"publisher","first-page":"e112102","DOI":"10.1371\/journal.pone.0112102","article-title":"Isolation of specific neurons from C. elegans Larvae for gene expression profiling","volume":"9","author":"Spencer","year":"2014","journal-title":"PLoS One"},{"key":"2026040114291463700_jkag030-B74","doi-asserted-by":"publisher","first-page":"iyae050","DOI":"10.1093\/genetics\/iyae050","article-title":"WormBase 2024: status and transitioning to Alliance infrastructure","volume":"227","author":"Sternberg","year":"2024","journal-title":"Genetics"},{"key":"2026040114291463700_jkag030-B75","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1186\/1471-2164-13-333","article-title":"A survey of putative secreted and transmembrane proteins encoded in the C. elegans genome","volume":"13","author":"Suh","year":"2012","journal-title":"BMC Genomics"},{"key":"2026040114291463700_jkag030-B76","doi-asserted-by":"publisher","first-page":"9279","DOI":"10.1523\/JNEUROSCI.6606-10.2011","article-title":"Spontaneous age-related neurite branching in Caenorhabditis elegans","volume":"31","author":"Tank","year":"2011","journal-title":"J Neurosci Off J Soc Neurosci"},{"key":"2026040114291463700_jkag030-B77","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1016\/j.cell.2013.07.006","article-title":"PQM-1 complements DAF-16 as a key transcriptional regulator of DAF-2-mediated development and longevity","volume":"154","author":"Tepper","year":"2013","journal-title":"Cell"},{"key":"2026040114291463700_jkag030-B78","doi-asserted-by":"publisher","first-page":"613","DOI":"10.1016\/j.neuron.2006.07.024","article-title":"The insulin\/PI 3-kinase pathway regulates salt chemotaxis learning in Caenorhabditis elegans","volume":"51","author":"Tomioka","year":"2006","journal-title":"Neuron"},{"key":"2026040114291463700_jkag030-B79","doi-asserted-by":"publisher","first-page":"11645","DOI":"10.1038\/ncomms11645","article-title":"Splicing factors control C. elegans behavioural learning in a single neuron by producing DAF-2c receptor","volume":"7","author":"Tomioka","year":"2016","journal-title":"Nat Commun"},{"key":"2026040114291463700_jkag030-B80","doi-asserted-by":"publisher","first-page":"8778","DOI":"10.1523\/JNEUROSCI.1494-11.2012","article-title":"Neurite sprouting and synapse deterioration in the aging Caenorhabditis elegans nervous system","volume":"32","author":"Toth","year":"2012","journal-title":"J Neurosci Off J Soc Neurosci"},{"key":"2026040114291463700_jkag030-B81","doi-asserted-by":"publisher","first-page":"RP95621","DOI":"10.7554\/eLife.95621","article-title":"The neuron-specific IIS\/FOXO transcriptome in aged animals reveals regulatory mechanisms of cognitive aging","volume":"13","author":"Weng","year":"2024","journal-title":"eLife"},{"key":"2026040114291463700_jkag030-B82","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1016\/j.cmet.2012.12.013","article-title":"Direct and indirect gene regulation by a life-extending FOXO protein in C. elegans: roles for GATA factors and lipid gene regulators","volume":"17","author":"Zhang","year":"2013","journal-title":"Cell Metab"},{"key":"2026040114291463700_jkag030-B83","doi-asserted-by":"publisher","first-page":"e13324","DOI":"10.1111\/acel.13324","article-title":"Mutation of daf-2 extends lifespan via tissue-specific effectors that suppress distinct life-limiting pathologies","volume":"20","author":"Zhao","year":"2021","journal-title":"Aging Cell"},{"key":"2026040114291463700_jkag030-B84","doi-asserted-by":"publisher","first-page":"112173","DOI":"10.1016\/j.mce.2024.112173","article-title":"C. elegans insulin-like peptides","volume":"585","author":"Zhu","year":"2024","journal-title":"Mol Cell Endocrinol"}],"container-title":["G3: Genes, Genomes, Genetics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/g3journal\/advance-article-pdf\/doi\/10.1093\/g3journal\/jkag030\/67204579\/jkag030.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/g3journal\/article-pdf\/16\/4\/jkag030\/67204579\/jkag030.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/g3journal\/article-pdf\/16\/4\/jkag030\/67204579\/jkag030.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T18:29:23Z","timestamp":1775068163000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/g3journal\/article\/doi\/10.1093\/g3journal\/jkag030\/8503865"}},"subtitle":[],"editor":[{"given":"M","family":"Zetka","sequence":"additional","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2026,3,3]]},"references-count":84,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2026,3,3]]},"published-print":{"date-parts":[[2026,4,1]]}},"URL":"https:\/\/doi.org\/10.1093\/g3journal\/jkag030","relation":{},"ISSN":["2160-1836"],"issn-type":[{"value":"2160-1836","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,4]]},"published":{"date-parts":[[2026,3,3]]},"article-number":"jkag030"}}