Fasting-based interventions are conserved regulators of aging that extend lifespan across species, including Caenorhabditis elegans. However, existing fasting studies are confounded by FUdR, antibiotics, germline-less mutants, and bacterial killing, which independently affect lifespan. FUdR alters lifespan, pathology, and stress responses, while antibiotics may affect worm physiology. To minimize these confounders, we developed an adult-onset intermittent fasting paradigm devoid of these factors. Wild-type worms subjected to daily fasting periods of 5 h, 6 h, or 18 h until day 10 of adulthood showed 24-57% lifespan extension. This tractable paradigm underscores the importance of limiting pharmacological and microbial conditions in dietary-intervention experiments.
","acknowledgements":"We thank all members of the Silva-García laboratory for their helpful discussions. Bristol N2 strain was provided by the Caenorhabditis Genetics Center (CGC), which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440).
","authors":[{"affiliations":["Brown University, Providence, RI, United States"],"departments":["Department of Molecular Biology, Cell Biology, and Biochemistry, Brown University. Providence, RI, USA ","Center on the Biology of Aging, Brown University. Providence, RI, USA ","Legorreta Cancer Center, Brown University. Providence, RI, USA ",""],"credit":["formalAnalysis","investigation","writing_originalDraft","dataCuration"],"email":"purbasha_dasgupta@brown.edu","firstName":"Purbasha","lastName":"Dasgupta","submittingAuthor":true,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0000-0001-6056-5722"},{"affiliations":["Brown University, Providence, RI, United States"],"departments":["Department of Molecular Biology, Cell Biology, and Biochemistry, Brown University. Providence, RI, USA ","Center on the Biology of Aging, Brown University. Providence, RI, USA ","Legorreta Cancer Center, Brown University. Providence, RI, USA "],"credit":["conceptualization","fundingAcquisition","writing_reviewEditing"],"email":"gio@brown.edu","firstName":"Carlos G.","lastName":"Silva-García","submittingAuthor":false,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"https://orcid.org/0000-0002-0434-7787"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":"https://portal.micropublication.org/uploads/05eb9881020281cfd960142986138667.csv"},"extendedData":[],"funding":"C.G.S-G. is funded by the National Institute on Aging R00AG065508 and National Institute of General Medical Sciences under Award Number P20GM156712 of the National Institutes of Health, the American Federation for Aging Research A24058, and Brown University (Division of Research Seed Award).
","image":{"url":"https://portal.micropublication.org/uploads/b85151c4b23c1e6ddab538fcfa27848a.png"},"imageCaption":"
(A–C) Schematics (top) and survival curves (bottom) of N2 wild-type worms under intermittent fasting (IF) regimens compared to ad libitum (AL) controls. Daily 5 h (A) and 6 h (B) fasting was initiated on day 1 of adulthood; 18 h overnight fasting (C) was initiated on day 3 after the reproductive peak. AL controls were maintained on seeded plates throughout. All regimens continued through day 10. No FUdR, antibiotics, or bacterial killing agents were used. Median lifespan was extended by 57% (5 h IF; p < 0.0001), 24% (6 h IF; p = 0.0011), and 35% (18 h IF; p < 0.0001) relative to AL controls (log-rank test). Table: Comparison of different fasting paradigms and their lifespan outcomes in C. elegans
","imageTitle":"Figure: Adult-onset intermittent fasting extends C. elegans lifespan independently of FUdR and antibiotics
","methods":"C. elegans strains and husbandry
Wild-type N2 animals were obtained from the Caenorhabditis Genetics Center (CGC). The strain was maintained on standard nematode growth media (NGM) seeded with E. coli OP50-1 at 20°C.
Bacterial strains
E. coli OP50-1 was cultured overnight in LB broth containing 50 μg/mL streptomycin at 37°C. For seeded plates, 100 μL of liquid culture was spread onto 60 mm NGM plates and allowed to grow for two days at room temperature prior to use.
Intermittent fasting regimen
Synchronized populations were obtained by bleaching gravid adults. Animals were raised on 60 mm seeded NGM plates without FUdR until day 1 of adulthood. Three daily fasting durations were tested: 5 h, and 6 h initiated at day 1 of adulthood, and 18 h initiated at day 3 (post-reproductive peak). Prior to each transfer, animals were individually picked onto an adjacent unseeded agar surface or a separate unseeded NGM plate and allowed to crawl briefly to shed residual bacteria from the cuticle. Then, animals were transferred on 100 mm unseeded NGM fasting plates containing no FUdR, antibiotics, or other bacterial killing agents. The 100 mm plate format was used to reduce animal loss from edge-crawling; plates were inspected for agar cracks prior to use. After each fasting interval, animals were transferred to freshly seeded 60 mm NGM plates using a small amount of bacteria from the destination plates as a cushion to minimize mechanical stress during transfer. This cycle was maintained through day 10 of adulthood. Ad libitum (AL) controls were maintained continuously on seeded plates without FUdR and subjected to identical handling to account for transfer effects.
Survival analyses
Lifespan assays were conducted at 20°C. Survival was scored daily through day 10, then every other day thereafter until death. Animals were considered dead upon failure to respond to three consecutive gentle prods with a platinum wire. Animals were censored for crawling off plates, bagging, vulval rupture, or plate contamination. To compensate for elevated censoring associated with intermittent fasting, cohort sizes were set at 300 worms for fasting conditions and 200 worms for ad libitum controls. Survival curves were compared using the log-rank (Mantel-Cox) method in GraphPad Prism.
","reagents":"","patternDescription":"
Dietary interventions are potent regulators of aging that can modulate lifespan across multiple species, including humans (Partridge et al., 2005; Kapahi et al., 2017; Green et al., 2022; Schmauck-Medina et al., 2026). Caenorhabditis elegans has emerged as a powerful model for such studies, owing to its short lifespan, conserved longevity pathways, and uniform single-bacterial diet, which permits precise nutritional manipulation while minimizing the behavioral and social complexities present in mammalian systems (Lee et al., 2006; Honjoh et al., 2009; Pang & Curran, 2014). Among dietary interventions, fasting-refeeding approaches, including intermittent fasting, temporary fasting, and time-restricted feeding, are particularly robust drivers of longevity through evolutionarily conserved mechanisms (Uno et al., 2013; de Cabo & Mattson, 2019; Ivimey-Cook et al., 2021; Longo et al. 2021; Tatge et al., 2026). However, standard fasting protocols in C. elegans frequently rely on experimental manipulations that independently influence lifespan and complicate the interpretation of dietary effects.
Prolonged food deprivation promotes matricidal hatching in reproductively active hermaphrodites, often necessitating treatment with 5-fluoro-2′-deoxyuridine FUdR. Yet FUdR independently alters stress resistance, age-associated pathology, and lifespan in a context-dependent manner and can interact with environmental stressors to further extend lifespan (Anderson et al., 2016; Wang et al., 2019). Because fasting is inherently a stress-responsive intervention, combining it with FUdR may obscure the specific contributions of fasting-induced mechanisms. Additional experimental variables arise from efforts to suppress bacterial proliferation on fasting plates. To prevent worms from seeding food-free agar with gut-retained or cuticle-bound E. coli, studies commonly employ bactericidal antibiotics (e.g., streptomycin or carbenicillin), UV- or heat-killed bacteria, or peptone-depleted media (Table 1). These approaches can independently affect longevity pathways. Antibiotics can extend lifespan by inducing mitonuclear protein imbalance and activating the mitochondrial unfolded protein response (UPRmt) (Houtkooper et al., 2013; Bonuccelli et al., 2023), whereas peptone depletion activates energy-sensing pathways, including AMPK and DAF-16/FoxO (Greer & Brunet, 2009; Stastna et al. 2015). Consequently, the lifespan extension observed under these conditions may reflect responses to pharmacologically or nutritional perturbations rather than physiological effects of fasting-refeeding cycles per se.
To minimize experimental confounders and build upon our previous studies of short-term fasting (Silva-Garcia et al., 2013; Lascarez-Lagunas et al., 2014; Huelgas-Morales et al., 2016), we developed an adult-onset intermittent fasting regimen that requires no FUdR, antibiotics, or other bacterial killing agents, on either fasting or fed plates. We tested three intermittent fasting protocols: 5 h and 6 h daily fasting initiated on day 1 of adulthood, and 18 h fasting initiated on day 3 of adulthood, after animals had reached their reproductive peak. All three regimens significantly extended median lifespan relative to ad libitum controls (5 h: 57% extension; 6 h: 24%; 18 h: 35%; log-rank p < 0.0001 for 5 h and 18 h, p = 0.0011 for 6 h; Figure 1A-C).
Together, these results demonstrate that short daily fasting bouts, as brief as 5 h, are sufficient to significantly extend lifespan under pharmacologically clean conditions. These findings are consistent with our previous work showing that short-term fasting profoundly remodels germline physiology by regulating apoptosis, stress granule formation, and protein translation (Silva-Garcia et al., 2013; Lascarez-Lagunas et al., 2014; Huelgas-Morales et al., 2016), highlighting the remarkable capacity of brief fasting periods to elicit broad physiological responses. Importantly, the longevity benefits observed in our study are comparable to those reported for established fasting-based interventions in C. elegans (Table 1).
Among the studies mentioned in Table 1, Li et al. (2026) is particularly noteworthy because its experimental design most closely resembles ours, employing short daily fasting windows in day 1 adults with live OP50 and reporting a modest but significant lifespan extension (25-34%). However, Li et al. acknowledge that the inclusion of FUdR limits their conclusions to relative lifespan comparisons performed under identical conditions. By contrast, our intermittent fasting paradigm is conducted entirely without FUdR, antibiotics, or other bacterial killing steps, allowing the longevity effects of fasting-refeeding cycles to be evaluated independently of pharmacological artifacts. Ongoing studies aim to define the molecular mechanisms underlying this response and to determine whether they overlap with or are distinct from those engaged by other fasting paradigms. Going forward, establishing the extent to which these mechanisms are conserved across species, including humans, will be critical for assessing their translational potential.
","references":[{"reference":"
Anderson EN, Corkins ME, Li JC, Singh K, Parsons S, Tucey TM, et al., Hart. 2016. C. elegans lifespan extension by osmotic stress requires FUdR, base excision repair, FOXO, and sirtuins. Mechanisms of Ageing and Development 154: 30-42.
","pubmedId":"","doi":"10.1016/j.mad.2016.01.004"},{"reference":"Bonuccelli G, Brooks DR, Shepherd S, Sotgia F, Lisanti MP. 2023. Antibiotics that target mitochondria extend lifespan in C. elegans. Aging 15: 11764-11781.
","pubmedId":"","doi":"10.18632/aging.205229"},{"reference":"de Cabo R, Mattson MP. 2019. Effects of Intermittent Fasting on Health, Aging, and Disease. New England Journal of Medicine 381: 2541-2551.
","pubmedId":"","doi":"10.1056/NEJMra1905136"},{"reference":"Green CL, Lamming DW, Fontana L. 2021. Molecular mechanisms of dietary restriction promoting health and longevity. Nature Reviews Molecular Cell Biology 23: 56-73.
","pubmedId":"","doi":"10.1038/s41580-021-00411-4"},{"reference":"Greer EL, Brunet A. 2009. Different dietary restriction regimens extend lifespan by both independent and overlapping genetic pathways in\n C. elegans. Aging Cell 8: 113-127.
","pubmedId":"","doi":"10.1111/j.1474-9726.2009.00459.x"},{"reference":"Honjoh S, Yamamoto T, Uno M, Nishida E. 2008. Signalling through RHEB-1 mediates intermittent fasting-induced longevity in C. elegans. Nature 457: 726-730.
","pubmedId":"","doi":"10.1038/nature07583"},{"reference":"Houtkooper RH, Mouchiroud L, Ryu D, Moullan N, Katsyuba E, Knott G, Williams RW, Auwerx J. 2013. Mitonuclear protein imbalance as a conserved longevity mechanism. Nature 497: 451-457.
","pubmedId":"","doi":"10.1038/nature12188"},{"reference":"Huelgas-Morales G, Silva-García CG, Salinas LS, Greenstein D, Navarro RE. 2016. The Stress Granule RNA-Binding Protein TIAR-1 Protects Female Germ Cells from Heat Shock inCaenorhabditis elegans. G3 Genes|Genomes|Genetics 6: 1031-1047.
","pubmedId":"","doi":"10.1534/g3.115.026815"},{"reference":"Ivimey-Cook ER, Sales K, Carlsson H, Immler S, Chapman T, Maklakov AA. 2021. Transgenerational fitness effects of lifespan extension by dietary restriction in\n Caenorhabditis elegans. Proceedings of the Royal Society B: Biological Sciences 288: 10.1098/rspb.2021.0701.
","pubmedId":"","doi":"10.1098/rspb.2021.0701"},{"reference":"Kaeberlein TL, Smith ED, Tsuchiya M, Welton KL, Thomas JH, Fields S, Kennedy BK, Kaeberlein M. 2006. Lifespan extension in\n Caenorhabditis elegans\n by complete removal of food. Aging Cell 5: 487-494.
","pubmedId":"","doi":"10.1111/j.1474-9726.2006.00238.x"},{"reference":"Kapahi P, Kaeberlein M, Hansen M. 2017. Dietary restriction and lifespan: Lessons from invertebrate models. Ageing Research Reviews 39: 3-14.
","pubmedId":"","doi":"10.1016/j.arr.2016.12.005"},{"reference":"Láscarez-Lagunas LI, Silva-García CG, Dinkova TD, Navarro RE. 2014. LIN-35/Rb Causes Starvation-Induced Germ Cell Apoptosis via CED-9/Bcl2 Downregulation in\n Caenorhabditis elegans. Molecular and Cellular Biology 34: 2499-2516.
","pubmedId":"","doi":"10.1128/MCB.01532-13"},{"reference":"Lee GD, Wilson MA, Zhu M, Wolkow CA, De Cabo R, Ingram DK, Zou S. 2006. Dietary deprivation extends lifespan in Caenorhabditis elegans. Aging Cell 5: 515-524.
","pubmedId":"","doi":"10.1111/j.1474-9726.2006.00241.x"},{"reference":"Li P, Zheng Y, Casas-Martinez JC, Xia Q, Miranda-Vizuete A, Goljanek-Whysall K, McDonagh B. 2026. Fasting requires Peroxiredoxin 2 and Peroxiredoxin 6 to coordinate redox dependent mitochondrial and lipid remodelling in Caenorhabditis elegans. Redox Biology 95: 104231.
","pubmedId":"","doi":"10.1016/j.redox.2026.104231"},{"reference":"Longo VD, Di Tano M, Mattson MP, Guidi N. 2021. Intermittent and periodic fasting, longevity and disease. Nature Aging 1: 47-59.
","pubmedId":"","doi":"10.1038/s43587-020-00013-3"},{"reference":"Pang S, Curran SP. 2014. Adaptive Capacity to Bacterial Diet Modulates Aging in C. elegans. Cell Metabolism 19: 221-231.
","pubmedId":"","doi":"10.1016/j.cmet.2013.12.005"},{"reference":"Partridge L, Piper MDW, Mair W. 2005. Dietary restriction in Drosophila. Mechanisms of Ageing and Development 126: 938-950.
","pubmedId":"","doi":"10.1016/j.mad.2005.03.023"},{"reference":"Schmauck-Medina T, Lautrup S, Di Francesco A, Mitchell SJ, Clemmensen C, Partridge L, et al., Fang. 2026. Dietary restriction in aging and longevity. Nature Aging 6: 485-505.
","pubmedId":"","doi":"10.1038/s43587-026-01091-5"},{"reference":"Carlos Giovanni SGa, Rosa EN. 2013. The\n C. elegans\n TIA‐1/TIAR homolog TIAR‐1 is required to induce germ cell apoptosis. genesis 51: 690-707.
","pubmedId":"","doi":"10.1002/dvg.22418"},{"reference":"Smith ED, Kaeberlein TL, Lydum BT, Sager J, Welton KL, Kennedy BK, Kaeberlein M. 2008. Age- and calorie-independent life span extension from dietary restriction by bacterial deprivation in Caenorhabditis elegans. BMC Developmental Biology 8: 10.1186/1471-213x-8-49.
","pubmedId":"","doi":"10.1186/1471-213X-8-49"},{"reference":"Stastna JJ, Snoek LB, Kammenga JE, Harvey SC. 2015. Genotype-dependent lifespan effects in peptone deprived Caenorhabditis elegans. Scientific Reports 5: 10.1038/srep16259.
","pubmedId":"","doi":"10.1038/srep16259"},{"reference":"Sutphin GL, Kaeberlein M. 2008. Dietary restriction by bacterial deprivation increases life span in wild-derived nematodes. Experimental Gerontology 43: 130-135.
","pubmedId":"","doi":"10.1016/j.exger.2007.10.019"},{"reference":"Tatge L, Kim J, Solano Fonseca R, Feola K, Wall JM, Otuzoglu G, et al., Douglas. 2026. Silencing lipid catabolism determines longevity in response to fasting. Nature Communications 17: 10.1038/s41467-026-68764-y.
","pubmedId":"","doi":"10.1038/s41467-026-68764-y"},{"reference":"Uno M, Honjoh S, Matsuda M, Hoshikawa H, Kishimoto S, Yamamoto T, et al., Nishida. 2013. A Fasting-Responsive Signaling Pathway that Extends Life Span in C. elegans. Cell Reports 3: 79-91.
","pubmedId":"","doi":"10.1016/j.celrep.2012.12.018"},{"reference":"Wang H, Zhao Y, Zhang Z. 2019. Age-dependent effects of floxuridine (FUdR) on senescent pathology and mortality in the nematode Caenorhabditis elegans. Biochemical and Biophysical Research Communications 509: 694-699.
","pubmedId":"","doi":"10.1016/j.bbrc.2018.12.161"},{"reference":"Zhou Y, Ahsan FM, Li S, Song M, Rotti JF, Yerevanian A, Shen Y, Soukas AA. 2026. Hormetic fasting extends Caenorhabditis elegans lifespan via H3K27 acetylation of lipid catabolism and antioxidant genes. Cell Reports 45: 117518.
","pubmedId":"","doi":"10.1016/j.celrep.2026.117518"}],"title":"A simplified intermittent fasting regimen robustly extends C. elegans lifespan without FUdR or antibiotic confounds
","reviews":[{"reviewer":{"displayName":"Teresa Lee"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":null}]},{"id":"a18ab7fe-c4a6-480b-9a47-5a9cfd25a621","decision":"accept","abstract":"Fasting-based interventions are conserved regulators of aging that extend lifespan across species, including Caenorhabditis elegans. However, existing fasting studies are confounded by FUdR, antibiotics, germline-less mutants, and bacterial killing, which independently affect lifespan. FUdR alters lifespan, pathology, and stress responses, while antibiotics may affect worm physiology. To minimize these confounders, we developed an adult-onset intermittent fasting paradigm devoid of these factors. Wild-type worms subjected to daily fasting periods of 5 h, 6 h, or 18 h until day 10 of adulthood showed 24-57% lifespan extension. This tractable paradigm underscores the importance of limiting pharmacological and microbial conditions in dietary-intervention experiments.
","acknowledgements":"We thank all members of the Silva-García laboratory for their helpful discussions. Bristol N2 strain was provided by the Caenorhabditis Genetics Center (CGC), which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440).
","authors":[{"affiliations":["Brown University, Providence, RI, United States"],"departments":["Department of Molecular Biology, Cell Biology, and Biochemistry, Brown University. Providence, RI, USA ","Center on the Biology of Aging, Brown University. Providence, RI, USA ","Legorreta Cancer Center, Brown University. Providence, RI, USA ",""],"credit":["formalAnalysis","investigation","writing_originalDraft","dataCuration"],"email":"purbasha_dasgupta@brown.edu","firstName":"Purbasha","lastName":"Dasgupta","submittingAuthor":true,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0000-0001-6056-5722"},{"affiliations":["Brown University, Providence, RI, United States"],"departments":["Department of Molecular Biology, Cell Biology, and Biochemistry, Brown University. Providence, RI, USA ","Center on the Biology of Aging, Brown University. Providence, RI, USA ","Legorreta Cancer Center, Brown University. Providence, RI, USA "],"credit":["conceptualization","fundingAcquisition","writing_reviewEditing"],"email":"gio@brown.edu","firstName":"Carlos G.","lastName":"Silva-García","submittingAuthor":false,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"https://orcid.org/0000-0002-0434-7787"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":"https://portal.micropublication.org/uploads/05eb9881020281cfd960142986138667.csv"},"extendedData":[],"funding":"C.G.S-G. is funded by the National Institute on Aging R00AG065508 and National Institute of General Medical Sciences under Award Number P20GM156712 of the National Institutes of Health, the American Federation for Aging Research A24058, and Brown University (Division of Research Seed Award).
","image":{"url":"https://portal.micropublication.org/uploads/b85151c4b23c1e6ddab538fcfa27848a.png"},"imageCaption":"
(A–C) Schematics (top) and survival curves (bottom) of N2 wild-type worms under intermittent fasting (IF) regimens compared to ad libitum (AL) controls. Daily 5 h (A) and 6 h (B) fasting was initiated on day 1 of adulthood; 18 h overnight fasting (C) was initiated on day 3 after the reproductive peak. AL controls were maintained on seeded plates throughout. All regimens continued through day 10. No FUdR, antibiotics, or bacterial killing agents were used. Median lifespan was extended by 57% (5 h IF; p < 0.0001), 24% (6 h IF; p = 0.0011), and 35% (18 h IF; p < 0.0001) relative to AL controls (log-rank test). Table: Comparison of different fasting paradigms and their lifespan outcomes in C. elegans
","imageTitle":"Figure: Adult-onset intermittent fasting extends C. elegans lifespan independently of FUdR and antibiotics
","methods":"C. elegans strains and husbandry
Wild-type N2 animals were obtained from the Caenorhabditis Genetics Center (CGC). The strain was maintained on standard nematode growth media (NGM) seeded with E. coli OP50-1 at 20°C.
Bacterial strains
E. coli OP50-1 was cultured overnight in LB broth containing 50 μg/mL streptomycin at 37°C. For seeded plates, 100 μL of liquid culture was spread onto 60 mm NGM plates and allowed to grow for two days at room temperature prior to use.
Intermittent fasting regimen
Synchronized populations were obtained by bleaching gravid adults. Animals were raised on 60 mm seeded NGM plates without FUdR until day 1 of adulthood. Three daily fasting durations were tested: 5 h, and 6 h initiated at day 1 of adulthood, and 18 h initiated at day 3 (post-reproductive peak). Prior to each transfer, animals were individually picked onto an adjacent unseeded agar surface or a separate unseeded NGM plate and allowed to crawl briefly to shed residual bacteria from the cuticle. Then, animals were transferred on 100 mm unseeded NGM fasting plates containing no FUdR, antibiotics, or other bacterial killing agents. The 100 mm plate format was used to reduce animal loss from edge-crawling; plates were inspected for agar cracks prior to use. After each fasting interval, animals were transferred to freshly seeded 60 mm NGM plates using a small amount of bacteria from the destination plates as a cushion to minimize mechanical stress during transfer. This cycle was maintained through day 10 of adulthood. Ad libitum (AL) controls were maintained continuously on seeded plates without FUdR and subjected to identical handling to account for transfer effects.
Survival analyses
Lifespan assays were conducted at 20°C. Survival was scored daily through day 10, then every other day thereafter until death. Animals were considered dead upon failure to respond to three consecutive gentle prods with a platinum wire. Animals were censored for crawling off plates, bagging, vulval rupture, or plate contamination. To compensate for elevated censoring associated with intermittent fasting, cohort sizes were set at 300 worms for fasting conditions and 200 worms for ad libitum controls. Survival curves were compared using the log-rank (Mantel-Cox) method in GraphPad Prism.
","reagents":"","patternDescription":"
Dietary interventions are potent regulators of aging that can modulate lifespan across multiple species, including humans (Partridge et al., 2005; Kapahi et al., 2017; Green et al., 2022; Schmauck-Medina et al., 2026). Caenorhabditis elegans has emerged as a powerful model for such studies, owing to its short lifespan, conserved longevity pathways, and uniform single-bacterial diet, which permits precise nutritional manipulation while minimizing the behavioral and social complexities present in mammalian systems (Lee et al., 2006; Honjoh et al., 2009; Pang & Curran, 2014). Among dietary interventions, fasting-refeeding approaches, including intermittent fasting, temporary fasting, and time-restricted feeding, are particularly robust drivers of longevity through evolutionarily conserved mechanisms (Uno et al., 2013; de Cabo & Mattson, 2019; Ivimey-Cook et al., 2021; Longo et al. 2021; Tatge et al., 2026). However, standard fasting protocols in C. elegans frequently rely on experimental manipulations that independently influence lifespan and complicate the interpretation of dietary effects.
Prolonged food deprivation promotes matricidal hatching in reproductively active hermaphrodites, often necessitating treatment with 5-fluoro-2′-deoxyuridine FUdR. Yet FUdR independently alters stress resistance, age-associated pathology, and lifespan in a context-dependent manner and can interact with environmental stressors to further extend lifespan (Anderson et al., 2016; Wang et al., 2019). Because fasting is inherently a stress-responsive intervention, combining it with FUdR may obscure the specific contributions of fasting-induced mechanisms. Additional experimental variables arise from efforts to suppress bacterial proliferation on fasting plates. To prevent worms from seeding food-free agar with gut-retained or cuticle-bound E. coli, studies commonly employ bactericidal antibiotics (e.g., streptomycin or carbenicillin), UV- or heat-killed bacteria, or peptone-depleted media (Table 1). These approaches can independently affect longevity pathways. Antibiotics can extend lifespan by inducing mitonuclear protein imbalance and activating the mitochondrial unfolded protein response (UPRmt) (Houtkooper et al., 2013; Bonuccelli et al., 2023), whereas peptone depletion activates energy-sensing pathways, including AMPK and DAF-16/FoxO (Greer & Brunet, 2009; Stastna et al. 2015). Consequently, the lifespan extension observed under these conditions may reflect responses to pharmacologically or nutritional perturbations rather than physiological effects of fasting-refeeding cycles per se.
To minimize experimental confounders and build upon our previous studies of short-term fasting (Silva-Garcia et al., 2013; Lascarez-Lagunas et al., 2014; Huelgas-Morales et al., 2016), we developed an adult-onset intermittent fasting regimen that requires no FUdR, antibiotics, or other bacterial killing agents, on either fasting or fed plates. We tested three intermittent fasting protocols: 5 h (Figure 1A) and 6 h (Figure 1B) daily fasting initiated on day 1 of adulthood, and 18 h fasting (Figure 1C) initiated on day 3 of adulthood, after animals had reached their reproductive peak. All three regimens significantly extended median lifespan relative to ad libitum controls (5 h: 57% extension; 6 h: 24%; 18 h: 35%; log-rank p < 0.0001 for 5 h and 18 h, p = 0.0011 for 6 h; Figure 1A-C).
Together, these results demonstrate that short daily fasting bouts, as brief as 5 h, are sufficient to significantly extend lifespan under pharmacologically clean conditions. These findings are consistent with our previous work showing that short-term fasting profoundly remodels germline physiology by regulating apoptosis, stress granule formation, and protein translation (Silva-Garcia et al., 2013; Lascarez-Lagunas et al., 2014; Huelgas-Morales et al., 2016), highlighting the remarkable capacity of brief fasting periods to elicit broad physiological responses. Importantly, the longevity benefits observed in our study are comparable to those reported for established fasting-based interventions in C. elegans (Table 1).
Among the studies mentioned in Table 1, Li et al. (2026) is particularly noteworthy because its experimental design most closely resembles ours, employing short daily fasting windows in day 1 adults with live OP50 and reporting a modest but significant lifespan extension (25-34%). However, Li et al. acknowledge that the inclusion of FUdR limits their conclusions to relative lifespan comparisons performed under identical conditions. By contrast, our intermittent fasting paradigm is conducted entirely without FUdR, antibiotics, or other bacterial killing steps, allowing the longevity effects of fasting-refeeding cycles to be evaluated independently of pharmacological artifacts. In our regimen, to minimize bacterial carryover onto fasting plates, animals were individually transferred to an unseeded agar surface and allowed to crawl briefly before transfer to fasting plates to reduce cuticle-associated bacteria. Shorter fasting intervals, particularly the 5 h and 6 h regimens, also limit the opportunity for bacterial proliferation compared with prolonged food deprivation. However, deposition and growth of gut-retained bacteria cannot be entirely excluded, particularly during the longer 18 h fasting interval. Ongoing studies aim to define the molecular mechanisms underlying this response and to determine whether they overlap with or are distinct from those engaged by other fasting paradigms. Going forward, establishing the extent to which these mechanisms are conserved across species, including humans, will be critical for assessing their translational potential.
","references":[{"reference":"
Anderson EN, Corkins ME, Li JC, Singh K, Parsons S, Tucey TM, et al., Hart. 2016. C. elegans lifespan extension by osmotic stress requires FUdR, base excision repair, FOXO, and sirtuins. Mechanisms of Ageing and Development 154: 30-42.
","pubmedId":"","doi":"10.1016/j.mad.2016.01.004"},{"reference":"Bonuccelli G, Brooks DR, Shepherd S, Sotgia F, Lisanti MP. 2023. Antibiotics that target mitochondria extend lifespan in C. elegans. Aging 15: 11764-11781.
","pubmedId":"","doi":"10.18632/aging.205229"},{"reference":"de Cabo R, Mattson MP. 2019. Effects of Intermittent Fasting on Health, Aging, and Disease. New England Journal of Medicine 381: 2541-2551.
","pubmedId":"","doi":"10.1056/NEJMra1905136"},{"reference":"Green CL, Lamming DW, Fontana L. 2021. Molecular mechanisms of dietary restriction promoting health and longevity. Nature Reviews Molecular Cell Biology 23: 56-73.
","pubmedId":"","doi":"10.1038/s41580-021-00411-4"},{"reference":"Greer EL, Brunet A. 2009. Different dietary restriction regimens extend lifespan by both independent and overlapping genetic pathways in\n C. elegans. Aging Cell 8: 113-127.
","pubmedId":"","doi":"10.1111/j.1474-9726.2009.00459.x"},{"reference":"Honjoh S, Yamamoto T, Uno M, Nishida E. 2008. Signalling through RHEB-1 mediates intermittent fasting-induced longevity in C. elegans. Nature 457: 726-730.
","pubmedId":"","doi":"10.1038/nature07583"},{"reference":"Houtkooper RH, Mouchiroud L, Ryu D, Moullan N, Katsyuba E, Knott G, Williams RW, Auwerx J. 2013. Mitonuclear protein imbalance as a conserved longevity mechanism. Nature 497: 451-457.
","pubmedId":"","doi":"10.1038/nature12188"},{"reference":"Huelgas-Morales G, Silva-García CG, Salinas LS, Greenstein D, Navarro RE. 2016. The Stress Granule RNA-Binding Protein TIAR-1 Protects Female Germ Cells from Heat Shock inCaenorhabditis elegans. G3 Genes|Genomes|Genetics 6: 1031-1047.
","pubmedId":"","doi":"10.1534/g3.115.026815"},{"reference":"Ivimey-Cook ER, Sales K, Carlsson H, Immler S, Chapman T, Maklakov AA. 2021. Transgenerational fitness effects of lifespan extension by dietary restriction in\n Caenorhabditis elegans. Proceedings of the Royal Society B: Biological Sciences 288: 10.1098/rspb.2021.0701.
","pubmedId":"","doi":"10.1098/rspb.2021.0701"},{"reference":"Kaeberlein TL, Smith ED, Tsuchiya M, Welton KL, Thomas JH, Fields S, Kennedy BK, Kaeberlein M. 2006. Lifespan extension in\n Caenorhabditis elegans\n by complete removal of food. Aging Cell 5: 487-494.
","pubmedId":"","doi":"10.1111/j.1474-9726.2006.00238.x"},{"reference":"Kapahi P, Kaeberlein M, Hansen M. 2017. Dietary restriction and lifespan: Lessons from invertebrate models. Ageing Research Reviews 39: 3-14.
","pubmedId":"","doi":"10.1016/j.arr.2016.12.005"},{"reference":"Láscarez-Lagunas LI, Silva-García CG, Dinkova TD, Navarro RE. 2014. LIN-35/Rb Causes Starvation-Induced Germ Cell Apoptosis via CED-9/Bcl2 Downregulation in\n Caenorhabditis elegans. Molecular and Cellular Biology 34: 2499-2516.
","pubmedId":"","doi":"10.1128/MCB.01532-13"},{"reference":"Lee GD, Wilson MA, Zhu M, Wolkow CA, De Cabo R, Ingram DK, Zou S. 2006. Dietary deprivation extends lifespan in Caenorhabditis elegans. Aging Cell 5: 515-524.
","pubmedId":"","doi":"10.1111/j.1474-9726.2006.00241.x"},{"reference":"Li P, Zheng Y, Casas-Martinez JC, Xia Q, Miranda-Vizuete A, Goljanek-Whysall K, McDonagh B. 2026. Fasting requires Peroxiredoxin 2 and Peroxiredoxin 6 to coordinate redox dependent mitochondrial and lipid remodelling in Caenorhabditis elegans. Redox Biology 95: 104231.
","pubmedId":"","doi":"10.1016/j.redox.2026.104231"},{"reference":"Longo VD, Di Tano M, Mattson MP, Guidi N. 2021. Intermittent and periodic fasting, longevity and disease. Nature Aging 1: 47-59.
","pubmedId":"","doi":"10.1038/s43587-020-00013-3"},{"reference":"Pang S, Curran SP. 2014. Adaptive Capacity to Bacterial Diet Modulates Aging in C. elegans. Cell Metabolism 19: 221-231.
","pubmedId":"","doi":"10.1016/j.cmet.2013.12.005"},{"reference":"Partridge L, Piper MDW, Mair W. 2005. Dietary restriction in Drosophila. Mechanisms of Ageing and Development 126: 938-950.
","pubmedId":"","doi":"10.1016/j.mad.2005.03.023"},{"reference":"Schmauck-Medina T, Lautrup S, Di Francesco A, Mitchell SJ, Clemmensen C, Partridge L, et al., Fang. 2026. Dietary restriction in aging and longevity. Nature Aging 6: 485-505.
","pubmedId":"","doi":"10.1038/s43587-026-01091-5"},{"reference":"Carlos Giovanni SGa, Rosa EN. 2013. The\n C. elegans\n TIA‐1/TIAR homolog TIAR‐1 is required to induce germ cell apoptosis. genesis 51: 690-707.
","pubmedId":"","doi":"10.1002/dvg.22418"},{"reference":"Smith ED, Kaeberlein TL, Lydum BT, Sager J, Welton KL, Kennedy BK, Kaeberlein M. 2008. Age- and calorie-independent life span extension from dietary restriction by bacterial deprivation in Caenorhabditis elegans. BMC Developmental Biology 8: 10.1186/1471-213x-8-49.
","pubmedId":"","doi":"10.1186/1471-213X-8-49"},{"reference":"Stastna JJ, Snoek LB, Kammenga JE, Harvey SC. 2015. Genotype-dependent lifespan effects in peptone deprived Caenorhabditis elegans. Scientific Reports 5: 10.1038/srep16259.
","pubmedId":"","doi":"10.1038/srep16259"},{"reference":"Sutphin GL, Kaeberlein M. 2008. Dietary restriction by bacterial deprivation increases life span in wild-derived nematodes. Experimental Gerontology 43: 130-135.
","pubmedId":"","doi":"10.1016/j.exger.2007.10.019"},{"reference":"Tatge L, Kim J, Solano Fonseca R, Feola K, Wall JM, Otuzoglu G, et al., Douglas. 2026. Silencing lipid catabolism determines longevity in response to fasting. Nature Communications 17: 10.1038/s41467-026-68764-y.
","pubmedId":"","doi":"10.1038/s41467-026-68764-y"},{"reference":"Uno M, Honjoh S, Matsuda M, Hoshikawa H, Kishimoto S, Yamamoto T, et al., Nishida. 2013. A Fasting-Responsive Signaling Pathway that Extends Life Span in C. elegans. Cell Reports 3: 79-91.
","pubmedId":"","doi":"10.1016/j.celrep.2012.12.018"},{"reference":"Wang H, Zhao Y, Zhang Z. 2019. Age-dependent effects of floxuridine (FUdR) on senescent pathology and mortality in the nematode Caenorhabditis elegans. Biochemical and Biophysical Research Communications 509: 694-699.
","pubmedId":"","doi":"10.1016/j.bbrc.2018.12.161"},{"reference":"Zhou Y, Ahsan FM, Li S, Song M, Rotti JF, Yerevanian A, Shen Y, Soukas AA. 2026. Hormetic fasting extends Caenorhabditis elegans lifespan via H3K27 acetylation of lipid catabolism and antioxidant genes. Cell Reports 45: 117518.
","pubmedId":"","doi":"10.1016/j.celrep.2026.117518"}],"title":"A simplified intermittent fasting regimen robustly extends C. elegans lifespan without FUdR or antibiotic confounds
","reviews":[{"reviewer":{"displayName":"Teresa Lee"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":"1791683589677"}]},{"id":"6f7878a2-e31e-4884-98dc-5ca7045a3a62","decision":"revise","abstract":"Fasting-based interventions are conserved regulators of aging that extend lifespan across species, including Caenorhabditis elegans. However, existing fasting studies are confounded by FUdR, antibiotics, germline-less mutants, and bacterial killing, which independently affect lifespan. FUdR alters lifespan, pathology, and stress responses, while antibiotics may affect worm physiology. To minimize these confounders, we developed an adult-onset intermittent fasting paradigm devoid of these factors. Wild-type worms subjected to daily fasting periods of 5 h, 6 h, or 18 h until day 10 of adulthood showed 24-57% lifespan extension. This tractable paradigm underscores the importance of limiting pharmacological and microbial conditions in dietary-intervention experiments.
","acknowledgements":"We thank all members of the Silva-García laboratory for their helpful discussions. Bristol N2 strain was provided by the Caenorhabditis Genetics Center (CGC), which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440).
","authors":[{"affiliations":["Brown University, Providence, RI, United States"],"departments":["Department of Molecular Biology, Cell Biology, and Biochemistry; Center on the Biology of Aging; Legorreta Cancer Center","","","","",""],"credit":["formalAnalysis","investigation","writing_originalDraft","dataCuration"],"email":"purbasha_dasgupta@brown.edu","firstName":"Purbasha","lastName":"Dasgupta","submittingAuthor":true,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0000-0001-6056-5722"},{"affiliations":["Brown University, Providence, RI, United States"],"departments":["Department of Molecular Biology, Cell Biology, and Biochemistry; Center on the Biology of Aging; Legorreta Cancer Center","",""],"credit":["conceptualization","fundingAcquisition","writing_reviewEditing"],"email":"gio@brown.edu","firstName":"Carlos G.","lastName":"Silva-García","submittingAuthor":false,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"https://orcid.org/0000-0002-0434-7787"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":"https://portal.micropublication.org/uploads/2989f9c1876bc88228876e2e07d67c68.csv"},"extendedData":[],"funding":"C.G.S-G. is funded by the National Institute on Aging R00AG065508 and National Institute of General Medical Sciences under Award Number P20GM156712 of the National Institutes of Health, the American Federation for Aging Research A24058, and Brown University (Division of Research Seed Award).
","image":{"url":"https://portal.micropublication.org/uploads/b85151c4b23c1e6ddab538fcfa27848a.png"},"imageCaption":"
(A–C) Schematics (top) and survival curves (bottom) of N2 wild-type worms under intermittent fasting (IF) regimens compared to ad libitum (AL) controls. Daily 5 h (A) and 6 h (B) fasting was initiated on day 1 of adulthood; 18 h overnight fasting (C) was initiated on day 3 after the reproductive peak. AL controls were maintained on seeded plates throughout. All regimens continued through day 10. No FUdR, antibiotics, or bacterial killing agents were used. Median lifespan was extended by 57% (5 h IF; p < 0.0001), 24% (6 h IF; p = 0.0011), and 35% (18 h IF; p < 0.0001) relative to AL controls (log-rank test). Table: Comparison of different fasting paradigms and their lifespan outcomes in C. elegans
","imageTitle":"Adult-onset intermittent fasting extends C. elegans lifespan independently of FUdR and antibiotics
","methods":"C. elegans strains and husbandry
Wild-type N2 animals were obtained from the Caenorhabditis Genetics Center (CGC). The strain was maintained on standard nematode growth media (NGM) seeded with E. coli OP50-1 at 20°C.
Bacterial strains
E. coli OP50-1 was cultured overnight in LB broth containing 50 μg/mL streptomycin at 37°C. For seeded plates, 100 μL of liquid culture was spread onto 60 mm NGM plates and allowed to grow for two days at room temperature prior to use.
Intermittent fasting regimen
Synchronized populations were obtained by bleaching gravid adults. Animals were raised on 60 mm seeded NGM plates without FUdR until day 1 of adulthood. Three daily fasting durations were tested: 5 h, and 6 h initiated on day 1 of adulthood, and 18 h initiated at day 3 (post-reproductive peak). Prior to each transfer, animals were individually picked onto an adjacent unseeded agar surface or a separate unseeded NGM plate and allowed to crawl briefly to shed residual bacteria from the cuticle. Then, animals were transferred on 100 mm unseeded NGM fasting plates containing no FUdR, antibiotics, or other bacterial killing agents. The 100 mm plate format was used to reduce animal loss from edge-crawling; plates were inspected for agar cracks prior to use. After each fasting interval, animals were transferred to freshly seeded 60 mm NGM plates using a small amount of bacteria from the destination plates as a cushion to minimize mechanical stress during transfer. This cycle was maintained through day 10 of adulthood. Ad libitum (AL) controls were maintained continuously on seeded plates without FUdR and subjected to identical handling to account for transfer effects.
Survival analyses
Lifespan assays were conducted at 20°C. Survival was scored daily through day 10, then every other day thereafter until death. Animals were considered dead upon failure to respond to three consecutive gentle prods with a platinum wire. Animals were censored for crawling off plates, bagging, vulval rupture, or plate contamination. To compensate for elevated censoring associated with intermittent fasting, cohort sizes were set at 300 worms for fasting conditions and 200 worms for ad libitum controls. Survival curves were compared using the log-rank (Mantel-Cox) method in GraphPad Prism.
","reagents":"","patternDescription":"
Dietary interventions are potent regulators of aging that can modulate lifespan across multiple species, including humans (Partridge et al., 2005; Kapahi et al., 2017; Green et al., 2021; Schmauck-Medina et al., 2026). Caenorhabditis elegans has emerged as a powerful model for such studies, owing to its short lifespan, conserved longevity pathways, and uniform single-bacterial diet, which permits precise nutritional manipulation while minimizing the behavioral and social complexities present in mammalian systems (Lee et al., 2006; Honjoh et al., 2009; Pang & Curran, 2014). Among dietary interventions, fasting-refeeding approaches, including intermittent fasting, temporary fasting, and time-restricted feeding, are particularly robust drivers of longevity through evolutionarily conserved mechanisms (Uno et al., 2013; de Cabo & Mattson, 2019; Ivimey-Cook et al., 2021; Longo et al., 2021; Tatge et al., 2026). However, standard fasting protocols in C. elegans frequently rely on experimental manipulations that independently influence lifespan and complicate the interpretation of dietary effects.
Prolonged food deprivation promotes matricidal hatching in reproductively active hermaphrodites, often necessitating treatment with 5-fluoro-2′-deoxyuridine (FUdR). Yet FUdR independently alters stress resistance, age-associated pathology, and lifespan in a context-dependent manner and can interact with environmental stressors to further extend lifespan (Anderson et al., 2016; Wang et al., 2019). Because fasting is inherently a stress-responsive intervention, combining it with FUdR may obscure the specific contributions of fasting-induced mechanisms. Additional experimental variables arise from efforts to suppress bacterial proliferation on fasting plates. To prevent worms from seeding food-free agar with gut-retained or cuticle-bound E. coli, studies commonly employ bactericidal antibiotics (e.g., streptomycin or carbenicillin), UV- or heat-killed bacteria, or peptone-depleted media (Table 1). These approaches can independently affect longevity pathways. Antibiotics can extend lifespan by inducing mitonuclear protein imbalance and activating the mitochondrial unfolded protein response (UPRmt) (Houtkooper et al., 2013; Bonuccelli et al., 2023), whereas peptone depletion activates energy-sensing pathways, including AMPK and DAF-16/FoxO (Greer & Brunet, 2009; Stastna et al., 2015). Consequently, the lifespan extension observed under these conditions may reflect responses to pharmacological or nutritional perturbations rather than physiological effects of fasting-refeeding cycles per se.
To minimize experimental confounders and build upon our previous studies of short-term fasting (Silva-García et al., 2013; Láscarez-Lagunas et al., 2014; Huelgas-Morales et al., 2016), we developed an adult-onset intermittent fasting regimen that requires no FUdR, antibiotics, or other bacterial killing agents, on either fasting or fed plates. We tested three intermittent fasting protocols: 5 h (Figure 1A) and 6 h (Figure 1B) daily fasting initiated on day 1 of adulthood, and 18 h fasting (Figure 1C) initiated on day 3 of adulthood, after animals had reached their reproductive peak. All three regimens significantly extended median lifespan relative to ad libitum controls (5 h: 57% extension; 6 h: 24%; 18 h: 35%; log-rank p < 0.0001 for 5 h and 18 h, p = 0.0011 for 6 h; Figure 1A-C).
Together, these results demonstrate that short daily fasting bouts, as brief as 5 h, are sufficient to significantly extend lifespan under pharmacologically clean conditions. These findings are consistent with our previous work showing that short-term fasting profoundly remodels germline physiology by regulating apoptosis, stress granule formation, and protein translation (Silva-García et al., 2013; Láscarez-Lagunas et al., 2014; Huelgas-Morales et al., 2016), highlighting the remarkable capacity of brief fasting periods to elicit broad physiological responses. Importantly, the longevity benefits observed in our study are comparable to those reported for established fasting-based interventions in C. elegans (Table 1).
Among the studies mentioned in Table 1, Li et al. (2026) is particularly noteworthy because its experimental design most closely resembles ours, employing short daily fasting windows in day 1 adults with live OP50 and reporting a modest but significant lifespan extension (25-34%). However, Li et al. acknowledge that the inclusion of FUdR limits their conclusions to relative lifespan comparisons performed under identical conditions. By contrast, our intermittent fasting paradigm is conducted entirely without FUdR, antibiotics, or other bacterial killing steps, allowing the longevity effects of fasting-refeeding cycles to be evaluated independently of pharmacological artifacts. In our regimen, to minimize bacterial carryover onto fasting plates, animals were individually transferred to an unseeded agar surface and allowed to crawl briefly before transfer to fasting plates to reduce cuticle-associated bacteria. Shorter fasting intervals, particularly the 5 h and 6 h regimens, also limit the opportunity for bacterial proliferation compared with prolonged food deprivation. However, deposition and growth of gut-retained bacteria cannot be entirely excluded, particularly during the longer 18 h fasting interval. Ongoing studies aim to define the molecular mechanisms underlying this response and to determine whether they overlap with or are distinct from those engaged by other fasting paradigms. Going forward, establishing the extent to which these mechanisms are conserved across species, including humans, will be critical for assessing their translational potential.
","references":[{"reference":"
Anderson EN, Corkins ME, Li JC, Singh K, Parsons S, Tucey TM, et al., Hart. 2016. C. elegans lifespan extension by osmotic stress requires FUdR, base excision repair, FOXO, and sirtuins. Mechanisms of Ageing and Development 154: 30-42.
","pubmedId":"","doi":"10.1016/j.mad.2016.01.004"},{"reference":"Bonuccelli G, Brooks DR, Shepherd S, Sotgia F, Lisanti MP. 2023. Antibiotics that target mitochondria extend lifespan in C. elegans. Aging 15: 11764-11781.
","pubmedId":"","doi":"10.18632/aging.205229"},{"reference":"de Cabo R, Mattson MP. 2019. Effects of Intermittent Fasting on Health, Aging, and Disease. New England Journal of Medicine 381: 2541-2551.
","pubmedId":"","doi":"10.1056/NEJMra1905136"},{"reference":"Green CL, Lamming DW, Fontana L. 2021. Molecular mechanisms of dietary restriction promoting health and longevity. Nature Reviews Molecular Cell Biology 23: 56-73.
","pubmedId":"","doi":"10.1038/s41580-021-00411-4"},{"reference":"Greer EL, Brunet A. 2009. Different dietary restriction regimens extend lifespan by both independent and overlapping genetic pathways in\n C. elegans. Aging Cell 8: 113-127.
","pubmedId":"","doi":"10.1111/j.1474-9726.2009.00459.x"},{"reference":"Honjoh S, Yamamoto T, Uno M, Nishida E. 2009. Signalling through RHEB-1 mediates intermittent fasting-induced longevity in C. elegans. Nature 457: 726-730.
","pubmedId":"","doi":"10.1038/nature07583"},{"reference":"Houtkooper RH, Mouchiroud L, Ryu D, Moullan N, Katsyuba E, Knott G, Williams RW, Auwerx J. 2013. Mitonuclear protein imbalance as a conserved longevity mechanism. Nature 497: 451-457.
","pubmedId":"","doi":"10.1038/nature12188"},{"reference":"Huelgas-Morales G, Silva-García CG, Salinas LS, Greenstein D, Navarro RE. 2016. The Stress Granule RNA-Binding Protein TIAR-1 Protects Female Germ Cells from Heat Shock in Caenorhabditis elegans. G3 Genes|Genomes|Genetics 6: 1031-1047.
","pubmedId":"","doi":"10.1534/g3.115.026815"},{"reference":"Ivimey-Cook ER, Sales K, Carlsson H, Immler S, Chapman T, Maklakov AA. 2021. Transgenerational fitness effects of lifespan extension by dietary restriction in\n Caenorhabditis elegans. Proceedings of the Royal Society B: Biological Sciences 288: 10.1098/rspb.2021.0701.
","pubmedId":"","doi":"10.1098/rspb.2021.0701"},{"reference":"Kaeberlein TL, Smith ED, Tsuchiya M, Welton KL, Thomas JH, Fields S, Kennedy BK, Kaeberlein M. 2006. Lifespan extension in\n Caenorhabditis elegans\n by complete removal of food. Aging Cell 5: 487-494.
","pubmedId":"","doi":"10.1111/j.1474-9726.2006.00238.x"},{"reference":"Kapahi P, Kaeberlein M, Hansen M. 2017. Dietary restriction and lifespan: Lessons from invertebrate models. Ageing Research Reviews 39: 3-14.
","pubmedId":"","doi":"10.1016/j.arr.2016.12.005"},{"reference":"Láscarez-Lagunas LI, Silva-García CG, Dinkova TD, Navarro RE. 2014. LIN-35/Rb Causes Starvation-Induced Germ Cell Apoptosis via CED-9/Bcl2 Downregulation in\n Caenorhabditis elegans. Molecular and Cellular Biology 34: 2499-2516.
","pubmedId":"","doi":"10.1128/MCB.01532-13"},{"reference":"Lee GD, Wilson MA, Zhu M, Wolkow CA, De Cabo R, Ingram DK, Zou S. 2006. Dietary deprivation extends lifespan in Caenorhabditis elegans. Aging Cell 5: 515-524.
","pubmedId":"","doi":"10.1111/j.1474-9726.2006.00241.x"},{"reference":"Li P, Zheng Y, Casas-Martinez JC, Xia Q, Miranda-Vizuete A, Goljanek-Whysall K, McDonagh B. 2026. Fasting requires Peroxiredoxin 2 and Peroxiredoxin 6 to coordinate redox dependent mitochondrial and lipid remodelling in Caenorhabditis elegans. Redox Biology 95: 104231.
","pubmedId":"","doi":"10.1016/j.redox.2026.104231"},{"reference":"Longo VD, Di Tano M, Mattson MP, Guidi N. 2021. Intermittent and periodic fasting, longevity and disease. Nature Aging 1: 47-59.
","pubmedId":"","doi":"10.1038/s43587-020-00013-3"},{"reference":"Pang S, Curran SP. 2014. Adaptive Capacity to Bacterial Diet Modulates Aging in C. elegans. Cell Metabolism 19: 221-231.
","pubmedId":"","doi":"10.1016/j.cmet.2013.12.005"},{"reference":"Partridge L, Piper MDW, Mair W. 2005. Dietary restriction in Drosophila. Mechanisms of Ageing and Development 126: 938-950.
","pubmedId":"","doi":"10.1016/j.mad.2005.03.023"},{"reference":"Schmauck-Medina T, Lautrup S, Di Francesco A, Mitchell SJ, Clemmensen C, Partridge L, et al., Fang. 2026. Dietary restriction in aging and longevity. Nature Aging 6: 485-505.
","pubmedId":"","doi":"10.1038/s43587-026-01091-5"},{"reference":"Silva-García CG, Rosa EN. 2013. The C. elegans TIA‐1/TIAR homolog TIAR‐1 is required to induce germ cell apoptosis. genesis 51: 690-707.
","pubmedId":"","doi":"10.1002/dvg.22418"},{"reference":"Smith ED, Kaeberlein TL, Lydum BT, Sager J, Welton KL, Kennedy BK, Kaeberlein M. 2008. Age- and calorie-independent life span extension from dietary restriction by bacterial deprivation in Caenorhabditis elegans. BMC Developmental Biology 8: 10.1186/1471-213x-8-49.
","pubmedId":"","doi":"10.1186/1471-213X-8-49"},{"reference":"Stastna JJ, Snoek LB, Kammenga JE, Harvey SC. 2015. Genotype-dependent lifespan effects in peptone deprived Caenorhabditis elegans. Scientific Reports 5: 10.1038/srep16259.
","pubmedId":"","doi":"10.1038/srep16259"},{"reference":"Sutphin GL, Kaeberlein M. 2008. Dietary restriction by bacterial deprivation increases life span in wild-derived nematodes. Experimental Gerontology 43: 130-135.
","pubmedId":"","doi":"10.1016/j.exger.2007.10.019"},{"reference":"Tatge L, Kim J, Solano Fonseca R, Feola K, Wall JM, Otuzoglu G, et al., Douglas. 2026. Silencing lipid catabolism determines longevity in response to fasting. Nature Communications 17: 10.1038/s41467-026-68764-y.
","pubmedId":"","doi":"10.1038/s41467-026-68764-y"},{"reference":"Uno M, Honjoh S, Matsuda M, Hoshikawa H, Kishimoto S, Yamamoto T, et al., Nishida. 2013. A Fasting-Responsive Signaling Pathway that Extends Life Span in C. elegans. Cell Reports 3: 79-91.
","pubmedId":"","doi":"10.1016/j.celrep.2012.12.018"},{"reference":"Wang H, Zhao Y, Zhang Z. 2019. Age-dependent effects of floxuridine (FUdR) on senescent pathology and mortality in the nematode Caenorhabditis elegans. Biochemical and Biophysical Research Communications 509: 694-699.
","pubmedId":"","doi":"10.1016/j.bbrc.2018.12.161"},{"reference":"Zhou Y, Ahsan FM, Li S, Song M, Rotti JF, Yerevanian A, Shen Y, Soukas AA. 2026. Hormetic fasting extends Caenorhabditis elegans lifespan via H3K27 acetylation of lipid catabolism and antioxidant genes. Cell Reports 45: 117518.
","pubmedId":"","doi":"10.1016/j.celrep.2026.117518"}],"title":"A simplified intermittent fasting regimen robustly extends C. elegans lifespan without FUdR or antibiotic confounds
","reviews":[],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":null}]},{"id":"ca2dec53-0327-4349-80ea-5b426be1b908","decision":"publish","abstract":"Fasting-based interventions are conserved regulators of aging that extend lifespan across species, including Caenorhabditis elegans. However, existing fasting studies are confounded by FUdR, antibiotics, germline-less mutants, and bacterial killing, which independently affect lifespan. FUdR alters lifespan, pathology, and stress responses, while antibiotics may affect worm physiology. To minimize these confounders, we developed an adult-onset intermittent fasting paradigm devoid of these factors. Wild-type worms subjected to daily fasting periods of 5 h, 6 h, or 18 h until day 10 of adulthood showed 24-57% lifespan extension. This tractable paradigm underscores the importance of limiting pharmacological and microbial conditions in dietary-intervention experiments.
","acknowledgements":"We thank all members of the Silva-García laboratory for their helpful discussions. Bristol N2 strain was provided by the Caenorhabditis Genetics Center (CGC), which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440).
","authors":[{"affiliations":["Brown University, Providence, RI, United States"],"departments":["Department of Molecular Biology, Cell Biology, and Biochemistry; Center on the Biology of Aging; Legorreta Cancer Center","","","","",""],"credit":["formalAnalysis","investigation","writing_originalDraft","dataCuration"],"email":"purbasha_dasgupta@brown.edu","firstName":"Purbasha","lastName":"Dasgupta","submittingAuthor":true,"correspondingAuthor":false,"equalContribution":false,"WBId":null,"orcid":"0000-0001-6056-5722"},{"affiliations":["Brown University, Providence, RI, United States"],"departments":["Department of Molecular Biology, Cell Biology, and Biochemistry; Center on the Biology of Aging; Legorreta Cancer Center","",""],"credit":["conceptualization","fundingAcquisition","writing_reviewEditing"],"email":"gio@brown.edu","firstName":"Carlos G.","lastName":"Silva-García","submittingAuthor":false,"correspondingAuthor":true,"equalContribution":false,"WBId":null,"orcid":"https://orcid.org/0000-0002-0434-7787"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":"https://portal.micropublication.org/uploads/2989f9c1876bc88228876e2e07d67c68.csv"},"extendedData":[],"funding":"C.G.S-G. is funded by the National Institute on Aging R00AG065508 and National Institute of General Medical Sciences under Award Number P20GM156712 of the National Institutes of Health, the American Federation for Aging Research A24058, and Brown University (Division of Research Seed Award).
","image":{"url":"https://portal.micropublication.org/uploads/b85151c4b23c1e6ddab538fcfa27848a.png"},"imageCaption":"
(A–C) Schematics (top) and survival curves (bottom) of N2 wild-type worms under intermittent fasting (IF) regimens compared to ad libitum (AL) controls. Daily 5 h (A) and 6 h (B) fasting was initiated on day 1 of adulthood; 18 h overnight fasting (C) was initiated on day 3 after the reproductive peak. AL controls were maintained on seeded plates throughout. All regimens continued through day 10. No FUdR, antibiotics, or bacterial killing agents were used. Median lifespan was extended by 57% (5 h IF; p < 0.0001), 24% (6 h IF; p = 0.0011), and 35% (18 h IF; p < 0.0001) relative to AL controls (log-rank test). Table: Comparison of different fasting paradigms and their lifespan outcomes in C. elegans
","imageTitle":"Adult-onset intermittent fasting extends C. elegans lifespan independently of FUdR and antibiotics
","methods":"C. elegans strains and husbandry
Wild-type N2 animals were obtained from the Caenorhabditis Genetics Center (CGC). The strain was maintained on standard nematode growth media (NGM) seeded with E. coli OP50-1 at 20°C.
Bacterial strains
E. coli OP50-1 was cultured overnight in LB broth containing 50 μg/mL streptomycin at 37°C. For seeded plates, 100 μL of liquid culture was spread onto 60 mm NGM plates and allowed to grow for two days at room temperature prior to use.
Intermittent fasting regimen
Synchronized populations were obtained by bleaching gravid adults. Animals were raised on 60 mm seeded NGM plates without FUdR until day 1 of adulthood. Three daily fasting durations were tested: 5 h, and 6 h initiated on day 1 of adulthood, and 18 h initiated at day 3 (post-reproductive peak). Prior to each transfer, animals were individually picked onto an adjacent unseeded agar surface or a separate unseeded NGM plate and allowed to crawl briefly to shed residual bacteria from the cuticle. Then, animals were transferred on 100 mm unseeded NGM fasting plates containing no FUdR, antibiotics, or other bacterial killing agents. The 100 mm plate format was used to reduce animal loss from edge-crawling; plates were inspected for agar cracks prior to use. After each fasting interval, animals were transferred to freshly seeded 60 mm NGM plates using a small amount of bacteria from the destination plates as a cushion to minimize mechanical stress during transfer. This cycle was maintained through day 10 of adulthood. Ad libitum (AL) controls were maintained continuously on seeded plates without FUdR and subjected to identical handling to account for transfer effects.
Survival analyses
Lifespan assays were conducted at 20°C. Survival was scored daily through day 10, then every other day thereafter until death. Animals were considered dead upon failure to respond to three consecutive gentle prods with a platinum wire. Animals were censored for crawling off plates, bagging, vulval rupture, or plate contamination. To compensate for elevated censoring associated with intermittent fasting, cohort sizes were set at 300 worms for fasting conditions and 200 worms for ad libitum controls. Survival curves were compared using the log-rank (Mantel-Cox) method in GraphPad Prism.
","reagents":"","patternDescription":"
Dietary interventions are potent regulators of aging that can modulate lifespan across multiple species, including humans (Partridge et al., 2005; Kapahi et al., 2017; Green et al., 2021; Schmauck-Medina et al., 2026). Caenorhabditis elegans has emerged as a powerful model for such studies, owing to its short lifespan, conserved longevity pathways, and uniform single-bacterial diet, which permits precise nutritional manipulation while minimizing the behavioral and social complexities present in mammalian systems (Lee et al., 2006; Honjoh et al., 2009; Pang & Curran, 2014). Among dietary interventions, fasting-refeeding approaches, including intermittent fasting, temporary fasting, and time-restricted feeding, are particularly robust drivers of longevity through evolutionarily conserved mechanisms (Uno et al., 2013; de Cabo & Mattson, 2019; Ivimey-Cook et al., 2021; Longo et al., 2021; Tatge et al., 2026). However, standard fasting protocols in C. elegans frequently rely on experimental manipulations that independently influence lifespan and complicate the interpretation of dietary effects.
Prolonged food deprivation promotes matricidal hatching in reproductively active hermaphrodites, often necessitating treatment with 5-fluoro-2′-deoxyuridine (FUdR). Yet FUdR independently alters stress resistance, age-associated pathology, and lifespan in a context-dependent manner and can interact with environmental stressors to further extend lifespan (Anderson et al., 2016; Wang et al., 2019). Because fasting is inherently a stress-responsive intervention, combining it with FUdR may obscure the specific contributions of fasting-induced mechanisms. Additional experimental variables arise from efforts to suppress bacterial proliferation on fasting plates. To prevent worms from seeding food-free agar with gut-retained or cuticle-bound E. coli, studies commonly employ bactericidal antibiotics (e.g., streptomycin or carbenicillin), UV- or heat-killed bacteria, or peptone-depleted media (Table 1). These approaches can independently affect longevity pathways. Antibiotics can extend lifespan by inducing mitonuclear protein imbalance and activating the mitochondrial unfolded protein response (UPRmt) (Houtkooper et al., 2013; Bonuccelli et al., 2023), whereas peptone depletion activates energy-sensing pathways, including AMPK and DAF-16/FoxO (Greer & Brunet, 2009; Stastna et al., 2015). Consequently, the lifespan extension observed under these conditions may reflect responses to pharmacological or nutritional perturbations rather than physiological effects of fasting-refeeding cycles per se.
To minimize experimental confounders and build upon our previous studies of short-term fasting (Silva-García et al., 2013; Láscarez-Lagunas et al., 2014; Huelgas-Morales et al., 2016), we developed an adult-onset intermittent fasting regimen that requires no FUdR, antibiotics, or other bacterial killing agents, on either fasting or fed plates. We tested three intermittent fasting protocols: 5 h (Figure 1A) and 6 h (Figure 1B) daily fasting initiated on day 1 of adulthood, and 18 h fasting (Figure 1C) initiated on day 3 of adulthood, after animals had reached their reproductive peak. All three regimens significantly extended median lifespan relative to ad libitum controls (5 h: 57% extension; 6 h: 24%; 18 h: 35%; log-rank p < 0.0001 for 5 h and 18 h, p = 0.0011 for 6 h; Figure 1A-C).
Together, these results demonstrate that short daily fasting bouts, as brief as 5 h, are sufficient to significantly extend lifespan under pharmacologically clean conditions. These findings are consistent with our previous work showing that short-term fasting profoundly remodels germline physiology by regulating apoptosis, stress granule formation, and protein translation (Silva-García et al., 2013; Láscarez-Lagunas et al., 2014; Huelgas-Morales et al., 2016), highlighting the remarkable capacity of brief fasting periods to elicit broad physiological responses. Importantly, the longevity benefits observed in our study are comparable to those reported for established fasting-based interventions in C. elegans (Table 1).
Among the studies mentioned in Table 1, Li et al. (2026) is particularly noteworthy because its experimental design most closely resembles ours, employing short daily fasting windows in day 1 adults with live OP50 and reporting a modest but significant lifespan extension (25-34%). However, Li et al. acknowledge that the inclusion of FUdR limits their conclusions to relative lifespan comparisons performed under identical conditions. By contrast, our intermittent fasting paradigm is conducted entirely without FUdR, antibiotics, or other bacterial killing steps, allowing the longevity effects of fasting-refeeding cycles to be evaluated independently of pharmacological artifacts. In our regimen, to minimize bacterial carryover onto fasting plates, animals were individually transferred to an unseeded agar surface and allowed to crawl briefly before transfer to fasting plates to reduce cuticle-associated bacteria. Shorter fasting intervals, particularly the 5 h and 6 h regimens, also limit the opportunity for bacterial proliferation compared with prolonged food deprivation. However, deposition and growth of gut-retained bacteria cannot be entirely excluded, particularly during the longer 18 h fasting interval. Ongoing studies aim to define the molecular mechanisms underlying this response and to determine whether they overlap with or are distinct from those engaged by other fasting paradigms. Going forward, establishing the extent to which these mechanisms are conserved across species, including humans, will be critical for assessing their translational potential.
","references":[{"reference":"
Anderson EN, Corkins ME, Li JC, Singh K, Parsons S, Tucey TM, et al., Hart. 2016. C. elegans lifespan extension by osmotic stress requires FUdR, base excision repair, FOXO, and sirtuins. Mechanisms of Ageing and Development 154: 30-42.
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","pubmedId":"","doi":"10.1016/j.celrep.2026.117518"}],"title":"A simplified intermittent fasting regimen robustly extends C. elegans lifespan without FUdR or antibiotic confounds
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adulthood","Experimental conditions":"FUdR (50 uM) on all plates; UV-killed OP50 for fed controls; N2 background","Effects on longevity/mortality":"~50% increase in median lifespan","References":"Kaeberlein et al. (2006)"},{"Fasting regimen":"Complete dietary deprivation (DD); initiated at day 2 of adulthood for optimal effect (also tested at days 4 and 8); worms remain food-deprived for the remainder of their lives","Experimental conditions":"FUdR (250 ug/mL), primarily fem-1(hc17) temperature-sensitive sterile mutant background","Effects on longevity/mortality":"Up to 42.5% mean lifespan extension (day 2 DD initiation in sterile adults); detrimental when imposed on reproductively active day 0 adults","References":"Lee et al. (2006)"},{"Fasting regimen":"Chronic bacterial deprivation (BD); complete removal of OP50 from NGM plates initiated at day 4 of adulthood; animals remain food-deprived for life. Also tested: BD initiated at days 8, 14, 20, and 24; and transient BD (day 2 to day 8, 14, or 28) with return to fed conditions","Experimental conditions":"FUdR (50 uM); UV-killed OP50 on control-fed plates; N2 background","Effects on longevity/mortality":"~48-67% median lifespan extension (day 4 BD onset, N2); significant extension even when BD initiated after >50% of cohort had died (day 24); transient BD conferred lasting survival and thermotolerance benefits after return to fed diet","References":"Smith et al. (2008)"},{"Fasting regimen":"Chronic bacterial deprivation from day 4 of adulthood","Experimental conditions":"UV-killed OP50 on control plates; FUdR 50 uM","Effects on longevity/mortality":"Mean LS +30%; max LS +30% (N2); wild isolates +23-51%","References":"Sutphin & Kaeberlein (2008)"},{"Fasting regimen":"Alternate-day fasting or every-2-day fasting","Experimental conditions":"FUdR (200 ug/mL), added at day 3 post-hatching, UV-killed OP50 was used for both AL controls and refeeding phases (not live bacteria); N2 background","Effects on longevity/mortality":"40.4% (alternate-day) and 56.6% (every-2-day) mean lifespan extension","References":"Honjoh et al. (2009)"},{"Fasting regimen":"2-day bacterial deprivation; late-L4 to day 2 of adulthood; returned to AL from day 3","Experimental conditions":"No FUdR, live OP50-1; no peptone on fasting plates; N2 background","Effects on longevity/mortality":"log-odds = -1.69 (reduced mortality)","References":"Ivimey-Cook et al. (2021)"},{"Fasting regimen":"Intermittent fasting: 4 h, 8 h or 12 h fasting/day for 5 consecutive days (days 1-5 of adulthood); refed on OP50 after each fast","Experimental conditions":"FUdR (10 uM, post-fasting lifespan plates), Live OP50; N2 background","Effects on longevity/mortality":"~27% (4 h); ~25% (8 h); ~34% (12 h) mean lifespan extension","References":"Li et al. (2026)"},{"Fasting regimen":"Single 24h fast at day 1 of adulthood; refed thereafter on standard NGM","Experimental conditions":"FUdR (50 mM as reported), live OP50-1; N2 background","Effects on longevity/mortality":"~11% - 67% median lifespan extension across replicates","References":"Zhou et al. (2026)"},{"Fasting regimen":"Single 24h fast at day 1 of adulthood; refed thereafter","Experimental conditions":"FUdR (100 ug/mL); carbenicillin (100 ug/mL); IPTG (1 mM); HT115 bacteria","Effects on longevity/mortality":"40.8% median lifespan extension","References":"Tatge et al. (2026)"},{"Fasting regimen":"5 h and 6 h daily fasting initiated on day 1 of adulthood; 18 h overnight fasting initiated on day 3 of adulthood (post attainment of reproductive peak); all regimens maintained through day 10.","Experimental conditions":"No FUdR, live OP50-1; no antibiotics or other bacteria-killing agents; N2 background","Effects on longevity/mortality":"57% (5 h IF), 24% (6 h IF), and 35% (18 h IF) median lifespan extension","References":"This study"}]}}}, "staticQueryHashes": ["2114697108"]}