{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T20:04:55Z","timestamp":1792785895379,"version":"4.5.0"},"reference-count":98,"publisher":"Oxford University Press (OUP)","license":[{"start":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T00:00:00Z","timestamp":1790294400000},"content-version":"am","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"name":"DBT-Wellcome Trust Intermediate Fellowship","award":["IA\/I\/20\/1\/504930"],"award-info":[{"award-number":["IA\/I\/20\/1\/504930"]}],"id":[{"id":"https:\/\/ror.org\/029chgv08","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001843","name":"Science and Engineering Research Board","doi-asserted-by":"publisher","award":["CRG\/2019\/002460"],"award-info":[{"award-number":["CRG\/2019\/002460"]}],"id":[{"id":"10.13039\/501100001843","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Insufficient protein intake causes malnutrition, compromising immunity and increasing the risk of pathogenic infections. In fact, under limited protein availability, organisms may experience stronger selective pressure to optimally allocate their resources between immunity vs other energy-demanding processes, such as reproduction, resulting in evolutionary trade-offs. In many species, protein deficiency may also more strongly affect female reproduction, leading to sexually dimorphic trade-offs involving immunity and infection outcomes, though these have not been experimentally tested. Here, we demonstrate that in\u00a0Drosophila melanogaster populations evolving greater investment in early reproduction in response to a lower-protein diet before oviposition, unmated females exhibited lower post-infection survival against Providencia rettgeri than unmated males, corroborating our prediction. However, mating eliminated these sex differences, with both sexes exhibiting nearly identical infection costs and reduced post-infection survival compared with their unselected counterparts. Overall, selected flies also had lower infection tolerance, although the effect on pathogen resistance was weak and inconsistent. Immune responses, including Toll- and IMD-mediated pathways, were either less upregulated or more downregulated in selected flies, which could explain their increased pathogen susceptibility. Downregulated metabolic pathways may also play a role. Overall, this study offers new evolutionary insights into how low-protein diets drive sex- and reproductive-status-specific variations in immunity and health.<\/jats:p>","DOI":"10.1093\/evolut\/qpag175","type":"journal-article","created":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T11:17:27Z","timestamp":1790335047000},"source":"Crossref","is-referenced-by-count":1,"title":["Adaptation to a lower protein diet before reproduction trades off with immunocompetence and the ability to survive pathogenic infections"],"prefix":"10.1093","author":[{"given":"Saubhik","family":"Sarkar","sequence":"first","affiliation":[{"name":"Trivedi School of Biosciences, Ashoka University , Sonipat, Haryana ,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dipendra Nath","family":"Basu","sequence":"additional","affiliation":[{"name":"Trivedi School of Biosciences, Ashoka University , Sonipat, Haryana ,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simran","family":"Sethi","sequence":"additional","affiliation":[{"name":"Trivedi School of Biosciences, Ashoka University , Sonipat, Haryana ,","place":["India"]},{"name":"Department of Biological Sciences, Indian Institute of Science Education and Research Berhampur , Ganjam, Odisha ,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9588-0316","authenticated-orcid":false,"given":"Bodhisatta","family":"Nandy","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Indian Institute of Science Education and Research Berhampur , Ganjam, Odisha ,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Imroze","family":"Khan","sequence":"additional","affiliation":[{"name":"Trivedi School of Biosciences, Ashoka University , Sonipat, Haryana ,","place":["India"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,9,25]]},"reference":[{"issue":"8","key":"2026101500270531600_bib1","doi-asserted-by":"publisher","first-page":"a017566","DOI":"10.1101\/cshperspect.a017566","article-title":"Sexual conflict, life span, and aging","volume":"6","author":"Adler","year":"2014","journal-title":"Cold Spring Harbor Perspectives in Biology"},{"issue":"6","key":"2026101500270531600_bib2","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1016\/j.exger.2013.03.007","article-title":"The lifespan-reproduction trade-off under dietary restriction is sex-specific and context-dependent","volume":"48","author":"Adler","year":"2013","journal-title":"Experimental Gerontology"},{"issue":"2","key":"2026101500270531600_bib3","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1093\/bioinformatics\/btu638","article-title":"HTSeq\u2014a Python framework to work with high-throughput sequencing data","volume":"31","author":"Anders","year":"2015","journal-title":"Bioinformatics"},{"issue":"1","key":"2026101500270531600_bib4","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/s0952-7915(99)00045-x","article-title":"Toll signaling pathways in the innate immune response","volume":"12","author":"Anderson","year":"2000","journal-title":"Current Opinion in Immunology"},{"key":"2026101500270531600_bib5","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1146\/annurev-immunol-020711-075030","article-title":"Tolerance of infections","volume":"30","author":"Ayres","year":"2012","journal-title":"Annual Review of Immunology"},{"key":"2026101500270531600_bib6","doi-asserted-by":"publisher","first-page":"3075","DOI":"10.3389\/fimmu.2019.03075","article-title":"Sexual dimorphisms in innate immunity and responses to infection in Drosophila melanogaster","volume":"10","author":"Belmonte","year":"2020","journal-title":"Frontiers in Immunology"},{"issue":"3","key":"2026101500270531600_bib7","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1111\/j.1365-2435.2008.01417.x","article-title":"Sexual selection, sexual conflict and the evolution of ageing and life span","volume":"22","author":"Bonduriansky","year":"2008","journal-title":"Functional Ecology"},{"issue":"2","key":"2026101500270531600_bib8","doi-asserted-by":"publisher","first-page":"378","DOI":"10.32614\/RJ-2017-066","article-title":"glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling","volume":"9","author":"Brooks","year":"2017","journal-title":"The R Journal"},{"issue":"1786","key":"2026101500270531600_bib9","doi-asserted-by":"publisher","first-page":"20140128","DOI":"10.1098\/rspb.2014.0128","article-title":"Protein-poor diet reduces host-specific immune gene expression in Bombus terrestris","volume":"281","author":"Brunner","year":"2014","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"11","key":"2026101500270531600_bib10","doi-asserted-by":"publisher","first-page":"5385","DOI":"10.1002\/ece3.4055","article-title":"Dietary choices are influenced by genotype, mating status, and sex in Drosophila melanogaster","volume":"8","author":"Camus","year":"2018","journal-title":"Ecology and Evolution"},{"issue":"1","key":"2026101500270531600_bib11","doi-asserted-by":"publisher","first-page":"iyab188","DOI":"10.1093\/genetics\/iyab188","article-title":"Cecropins contribute to Drosophila host defense against a subset of fungal and Gram-negative bacterial infection","volume":"220","author":"Carboni","year":"2022","journal-title":"Genetics"},{"key":"2026101500270531600_bib12","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1002\/9780470376980","article-title":"Nutrition and nutritional problems in wild animal. FOWLER, ME; CUBAS, ZS biology, medicine, and surgery of south American wild animals","author":"Carciofi","year":"2001"},{"issue":"3","key":"2026101500270531600_bib13","doi-asserted-by":"publisher","first-page":"564","DOI":"10.1111\/j.1420-9101.2008.01664.x","article-title":"Mating triggers dynamic immune regulations in wood ant queens","volume":"22","author":"Castella","year":"2009","journal-title":"Journal of Evolutionary Biology"},{"issue":"1940","key":"2026101500270531600_bib14","doi-asserted-by":"publisher","first-page":"20202684","DOI":"10.1098\/rspb.2020.2684","article-title":"Experimental evolution of post-ingestive nutritional compensation in response to a nutrient-poor diet","volume":"287","author":"Cavigliasso","year":"2020","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"10","key":"2026101500270531600_bib15","doi-asserted-by":"publisher","first-page":"e0224440","DOI":"10.1371\/journal.pone.0224440","article-title":"Consequences of chronic bacterial infection in Drosophila melanogaster","volume":"14","author":"Chambers","year":"2019","journal-title":"PLoS ONE"},{"issue":"8","key":"2026101500270531600_bib16","doi-asserted-by":"publisher","first-page":"e71984","DOI":"10.1371\/journal.pone.0071984","article-title":"BmPAH catalyzes the initial melanin biosynthetic step in Bombyx mori","volume":"8","author":"Chen","year":"2013","journal-title":"PLoS ONE"},{"issue":"9","key":"2026101500270531600_bib17","doi-asserted-by":"publisher","first-page":"e9287","DOI":"10.1002\/ece3.9287","article-title":"Does perception of female cues modulate male short-term fitness components in Drosophila melanogaster?","volume":"12","author":"Corbel","year":"2022","journal-title":"Ecology and Evolution"},{"issue":"1","key":"2026101500270531600_bib18","doi-asserted-by":"publisher","first-page":"186","DOI":"10.1111\/j.1365-2435.2010.01766.x","article-title":"Macronutrient balance mediates trade-offs between immune function and life history traits","volume":"25","author":"Cotter","year":"2011","journal-title":"Functional Ecology"},{"issue":"8","key":"2026101500270531600_bib19","doi-asserted-by":"publisher","first-page":"1836","DOI":"10.1111\/evo.14560","article-title":"Evolution of a novel female reproductive strategy in Drosophila melanogaster populations subjected to long-term protein restriction","volume":"76","author":"Dasgupta","year":"2022","journal-title":"Evolution; Internation Journal of Organic Evolution"},{"key":"2026101500270531600_bib20","doi-asserted-by":"publisher","DOI":"10.1101\/2024.08.24.609539","article-title":"Adaptation to yeast restriction through the evolution of resource acquisition, macronutrient reserve, and ovarian function","volume-title":"bioRxiv","author":"Dasgupta","year":"2024"},{"key":"2026101500270531600_bib21","first-page":"259","article-title":"The energetics of immunity","volume":"1","author":"Demas","year":"2012","journal-title":"Ecoimmunology"},{"issue":"1","key":"2026101500270531600_bib22","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1016\/j.immuni.2017.06.016","article-title":"Peroxisome-mediated metabolism is required for immune response to microbial infection","volume":"47","author":"Di\u00a0Cara","year":"2017","journal-title":"Immunity"},{"issue":"5","key":"2026101500270531600_bib23","doi-asserted-by":"publisher","first-page":"3793","DOI":"10.1111\/pce.15361","article-title":"Multiple, single trait GWAS and supervised machine learning reveal the genetic architecture of Fraxinus excelsior tolerance to ash dieback in Europe","volume":"48","author":"Doonan","year":"2025","journal-title":"Plant, Cell & Environment"},{"issue":"1","key":"2026101500270531600_bib24","doi-asserted-by":"publisher","first-page":"2453","DOI":"10.1038\/s41467-021-22712-0","article-title":"Sight of parasitoid wasps accelerates sexual behavior and upregulates a micropeptide gene in Drosophila","volume":"12","author":"Ebrahim","year":"2021","journal-title":"Nature Communications"},{"issue":"1614","key":"2026101500270531600_bib25","doi-asserted-by":"publisher","first-page":"1211","DOI":"10.1098\/rspb.2006.0394","article-title":"Post-mating disparity between potential and realized immune response in Drosophila melanogaster","volume":"274","author":"Fedorka","year":"2007","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"11","key":"2026101500270531600_bib26","doi-asserted-by":"publisher","first-page":"2478","DOI":"10.1111\/j.0014-3820.2004.tb00877.x","article-title":"Immune suppression and the cost of reproduction in the ground cricket, Allonemobius socius","volume":"58","author":"Fedorka","year":"2004","journal-title":"Evolution; Internation Journal of Organic Evolution"},{"key":"2026101500270531600_bib27","doi-asserted-by":"publisher","DOI":"10.1093\/acprof:oso\/9780199568765.001.0001","volume-title":"Mechanisms of life history evolution: The genetics and physiology of life history traits and trade-offs","author":"Flatt","year":"2011"},{"issue":"3","key":"2026101500270531600_bib28","doi-asserted-by":"publisher","first-page":"515","DOI":"10.1093\/biomet\/81.3.515","article-title":"Proportional hazards tests and diagnostics based on weighted residuals","volume":"81","author":"Grambsch","year":"1994","journal-title":"Biometrika"},{"issue":"7","key":"2026101500270531600_bib29","doi-asserted-by":"publisher","first-page":"170342","DOI":"10.1098\/rsos.170342","article-title":"Nonlinear disease tolerance curves reveal distinct components of host responses to viral infection","volume":"4","author":"Gupta","year":"2017","journal-title":"Royal Society Open Science"},{"key":"2026101500270531600_bib30","doi-asserted-by":"publisher","first-page":"e44341","DOI":"10.7554\/eLife.44341","article-title":"Synergy and remarkable specificity of antimicrobial peptides in vivo using a systematic knockout approach","volume":"8","author":"Hanson","year":"2019","journal-title":"eLife"},{"issue":"6","key":"2026101500270531600_bib31","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41559-017-0152","article-title":"Perceptive costs of reproduction drive ageing and physiology in male Drosophila","volume":"1","author":"Harvanek","year":"2017","journal-title":"Nature Ecology & Evolution"},{"issue":"6","key":"2026101500270531600_bib32","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1177\/1535370218758275","article-title":"Impacts of maternal dietary protein intake on fetal survival, growth, and development","volume":"243","author":"Herring","year":"2018","journal-title":"Experimental Biology and Medicine"},{"issue":"3","key":"2026101500270531600_bib33","doi-asserted-by":"publisher","first-page":"445","DOI":"10.1101\/gr.185579.114","article-title":"The Release 6 reference sequence of the Drosophila melanogaster genome","volume":"25","author":"Hoskins","year":"2015","journal-title":"Genome Research"},{"issue":"6","key":"2026101500270531600_bib34","doi-asserted-by":"publisher","first-page":"4149","DOI":"10.1016\/j.ygeno.2021.10.018","article-title":"RNA-seq library preparation for comprehensive transcriptome analysis in cancer cells: The impact of insert size","volume":"113","author":"Jaksik","year":"2021","journal-title":"Genomics"},{"issue":"4","key":"2026101500270531600_bib35","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1111\/acel.12333","article-title":"Sex-specific effects of protein and carbohydrate intake on reproduction but not lifespan in Drosophila melanogaster","volume":"14","author":"Jensen","year":"2015","journal-title":"Aging Cell"},{"key":"2026101500270531600_bib36","volume-title":"survminer: Drawing survival curves using \u2018ggplot2\u2019","author":"Kassambara","year":"2016"},{"issue":"3\u20134","key":"2026101500270531600_bib37","doi-asserted-by":"publisher","first-page":"290","DOI":"10.1111\/j.1439-0426.1995.tb00029.x","article-title":"Protein and energy needs for maintenance and growth of Nile tilapia (Oreochromis niloticus)","volume":"11","author":"Kaushik","year":"1995","journal-title":"Journal of Applied Ichthyology"},{"issue":"2","key":"2026101500270531600_bib38","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1016\/j.cmet.2020.06.010","article-title":"Amino assets: How amino acids support immunity","volume":"32","author":"Kelly","year":"2020","journal-title":"Cell Metabolism"},{"issue":"1850","key":"2026101500270531600_bib39","doi-asserted-by":"publisher","first-page":"20170125","DOI":"10.1098\/rspb.2017.0125","article-title":"Early-life inflammation, immune response and ageing","volume":"284","author":"Khan","year":"2017","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"key":"2026101500270531600_bib40","first-page":"79","article-title":"Sex-specific effect of bacterial infection on components of adult fitness in Drosophila melanogaster","volume":"3","author":"Khan","year":"2011","journal-title":"Journal of Evolutionary Biology Research"},{"key":"2026101500270531600_bib41","article-title":"pheatmap: Pretty heatmaps","author":"Kolde","year":"2025"},{"issue":"4","key":"2026101500270531600_bib42","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1038\/sj.hdy.6801092","article-title":"Selection for resistance to a fungal pathogen in Drosophila melanogaster","volume":"100","author":"Kraaijeveld","year":"2008","journal-title":"Heredity"},{"key":"2026101500270531600_bib43","unstructured":"Krassowski M. (2020) ComplexUpset. Zenodo. 10.5281\/zenodo.3700590"},{"issue":"6","key":"2026101500270531600_bib44","doi-asserted-by":"publisher","first-page":"bio042952","DOI":"10.1242\/bio.042952","article-title":"Evidence of dietary protein restriction regulating pupation height, development time and lifespan in Drosophila melanogaster","volume":"8","author":"Krittika","year":"2019","journal-title":"Biology Open"},{"issue":"11","key":"2026101500270531600_bib45","doi-asserted-by":"publisher","first-page":"695","DOI":"10.1016\/j.exger.2009.08.001","article-title":"Contrasted effects of suppressing live yeast from food on longevity, aging and resistance to several stresses in Drosophila melanogaster","volume":"44","author":"Le\u00a0Rohellec","year":"2009","journal-title":"Experimental Gerontology"},{"key":"2026101500270531600_bib46","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/j.jinsphys.2015.02.007","article-title":"Dietary protein:carbohydrate balance is a critical modulator of lifespan and reproduction in Drosophila melanogaster: A test using a chemically defined diet","volume":"75","author":"Lee","year":"2015","journal-title":"Journal of Insect Physiology"},{"issue":"5","key":"2026101500270531600_bib47","doi-asserted-by":"publisher","first-page":"987","DOI":"10.1016\/j.anbehav.2013.08.018","article-title":"Sexual dimorphism in nutrient intake and life span is mediated by mating in Drosophila melanogaster","volume":"86","author":"Lee","year":"2013","journal-title":"Animal Behaviour"},{"issue":"6","key":"2026101500270531600_bib48","doi-asserted-by":"publisher","first-page":"1052","DOI":"10.1111\/j.1365-2435.2008.01459.x","article-title":"Dietary protein-quality influences melanization and immune function in an insect","volume":"22","author":"Lee","year":"2008","journal-title":"Functional Ecology"},{"issue":"6","key":"2026101500270531600_bib49","doi-asserted-by":"publisher","first-page":"973","DOI":"10.1016\/s0092-8674(00)80172-5","article-title":"The dorsoventral regulatory gene cassette sp\u00e4tzle\/Toll\/cactus controls the potent antifungal response in Drosophila adults","volume":"86","author":"Lemaitre","year":"1996","journal-title":"Cell"},{"issue":"12","key":"2026101500270531600_bib50","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biology"},{"issue":"1-2","key":"2026101500270531600_bib51","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1016\/s0166-4328(00)00164-9","article-title":"Behavioral effects of protein deprivation and rehabilitation in adult rats: Relevance to morphological alterations in the hippocampal formation","volume":"112","author":"Lukoyanov","year":"2000","journal-title":"Behavioural Brain Research"},{"key":"2026101500270531600_bib52","doi-asserted-by":"publisher","first-page":"104050","DOI":"10.1016\/j.jinsphys.2020.104050","article-title":"Amino acid quality modifies the quantitative availability of protein for reproduction in Drosophila melanogaster","volume":"139","author":"Ma","year":"2022","journal-title":"Journal of Insect Physiology"},{"key":"2026101500270531600_bib53","doi-asserted-by":"publisher","first-page":"107231","DOI":"10.1016\/j.vascn.2022.107231","article-title":"Protein malnutrition in BALB\/C mice: A model mimicking clinical scenario of marasmic-kwashiorkor malnutrition","volume":"119","author":"Madhura","year":"2023","journal-title":"Journal of Pharmacological and Toxicological Methods"},{"issue":"14","key":"2026101500270531600_bib54","doi-asserted-by":"publisher","first-page":"1062","DOI":"10.1016\/j.cub.2008.06.059","article-title":"Sex-specific fitness effects of nutrient intake on reproduction and lifespan","volume":"18","author":"Maklakov","year":"2008","journal-title":"Current Biology"},{"issue":"11","key":"2026101500270531600_bib55","doi-asserted-by":"publisher","first-page":"e1009192","DOI":"10.1371\/journal.pgen.1009192","article-title":"Innate immune signaling in Drosophila shifts anabolic lipid metabolism from triglyceride storage to phospholipid synthesis to support immune function","volume":"16","author":"Mart\u00ednez","year":"2020","journal-title":"PLOS Genetics"},{"issue":"7","key":"2026101500270531600_bib56","doi-asserted-by":"publisher","first-page":"1510","DOI":"10.1111\/j.0014-3820.2005.tb01800.x","article-title":"Bateman\u2019s principle and immunity: Phenotypically plastic reproductive strategies predict changes in immunological sex differences","volume":"59","author":"McKean","year":"2005","journal-title":"Evolution; Internation Journal of Organic Evolution"},{"issue":"2","key":"2026101500270531600_bib57","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1111\/j.1558-5646.2007.00286.x","article-title":"Sexual selection and immune function in Drosophila melanogaster","volume":"62","author":"McKean","year":"2008","journal-title":"Evolution; Internation Journal of Organic Evolution"},{"issue":"5","key":"2026101500270531600_bib58","doi-asserted-by":"publisher","first-page":"e8960","DOI":"10.1002\/ece3.8960","article-title":"Metabolic and immunological responses of Drosophila melanogaster to dietary restriction and bacterial infection differ substantially between genotypes in a population","volume":"12","author":"Meshrif","year":"2022","journal-title":"Ecology and Evolution"},{"key":"2026101500270531600_bib59","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/j.cois.2018.08.006","article-title":"Turning food into eggs: Insights from nutritional biology and developmental physiology of Drosophila","volume":"31","author":"Mirth","year":"2019","journal-title":"Current Opinion in Insect Science"},{"issue":"11","key":"2026101500270531600_bib60","doi-asserted-by":"publisher","first-page":"108506","DOI":"10.1016\/j.celrep.2020.108506","article-title":"Evidence for long-lasting transgenerational antiviral immunity in insects","volume":"33","author":"Mondotte","year":"2020","journal-title":"Cell Reports"},{"issue":"5494","key":"2026101500270531600_bib61","doi-asserted-by":"publisher","first-page":"1166","DOI":"10.1126\/science.290.5494.1166","article-title":"Survival for immunity: The price of immune system activation for bumblebee workers","volume":"290","author":"Moret","year":"2000","journal-title":"Science"},{"issue":"1-3","key":"2026101500270531600_bib62","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1016\/j.bbagen.2003.10.004","article-title":"Antibacterial activity and mechanism of action of tick defensin against Gram-positive bacteria","volume":"1624","author":"Nakajima","year":"2003","journal-title":"Biochimica et Biophysica Acta (BBA)\u2014General Subjects"},{"issue":"6","key":"2026101500270531600_bib63","doi-asserted-by":"publisher","first-page":"e0130334","DOI":"10.1371\/journal.pone.0130334","article-title":"The role of storage lipids in the relation between fecundity, locomotor activity, and lifespan of Drosophila melanogaster longevity-selected and control lines","volume":"10","author":"Nasiri\u00a0Moghadam","year":"2015","journal-title":"PLoS ONE"},{"key":"2026101500270531600_bib64","volume-title":"RColorBrewer: Colorbrewer palettes","author":"Neuwirth","year":"2022"},{"key":"2026101500270531600_bib65","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1163\/15707563-20191082","article-title":"Does mating negatively affect female immune defences in insects?","volume":"69","author":"Oku","year":"2019","journal-title":"Animal Biology"},{"issue":"2 Suppl","key":"2026101500270531600_bib66","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1016\/j.taap.2005.01.060","article-title":"How do male germ cells handle DNA damage?","volume":"207","author":"Olsen","year":"2005","journal-title":"Toxicology and Applied Pharmacology"},{"issue":"2","key":"2026101500270531600_bib67","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1111\/j.1365-2656.2008.01499.x","article-title":"Can the protein costs of bacterial resistance be offset by altered feeding behaviour?","volume":"78","author":"Povey","year":"2009","journal-title":"Journal of Animal Ecology"},{"issue":"1","key":"2026101500270531600_bib68","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1111\/1365-2656.12127","article-title":"Dynamics of macronutrient self-medication and illness-induced anorexia in virally infected insects","volume":"83","author":"Povey","year":"2014","journal-title":"Journal of Animal Ecology"},{"key":"2026101500270531600_bib69","first-page":"208","article-title":"Dietary protein requirement for maintenance, growth, and reproduction in fish: A review","volume":"8","author":"Radhakrishnan","year":"2020","journal-title":"J Entomol Zool Stud"},{"issue":"12","key":"2026101500270531600_bib70","doi-asserted-by":"publisher","first-page":"e238","DOI":"10.1371\/journal.pgen.0030238","article-title":"Sustained post-mating response in Drosophila melanogaster requires multiple seminal fluid proteins","volume":"3","author":"Ram","year":"2007","journal-title":"PLoS Genetics"},{"issue":"1","key":"2026101500270531600_bib71","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1086\/670649","article-title":"Sex-specific fitness consequences of nutrient intake and the evolvability of diet preferences","volume":"182","author":"Reddiex","year":"2013","journal-title":"The American Naturalist"},{"issue":"6","key":"2026101500270531600_bib72","doi-asserted-by":"publisher","first-page":"644","DOI":"10.1002\/evl3.261","article-title":"Transcriptomic evidence for a trade-off between germline proliferation and immunity in Drosophila","volume":"5","author":"Rodrigues","year":"2021","journal-title":"Evolution Letters"},{"issue":"1493","key":"2026101500270531600_bib73","doi-asserted-by":"publisher","first-page":"867","DOI":"10.1098\/rspb.2002.1959","article-title":"Bateman\u2019s principle and immunity","volume":"269","author":"Rolff","year":"2002","journal-title":"Proceedings of the Royal Society of London. Series B: Biological Sciences"},{"issue":"15","key":"2026101500270531600_bib74","doi-asserted-by":"publisher","first-page":"9916","DOI":"10.1073\/pnas.152271999","article-title":"Copulation corrupts immunity: A mechanism for a cost of mating in insects","volume":"99","author":"Rolff","year":"2002","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2026101500270531600_bib75","doi-asserted-by":"publisher","first-page":"11783","DOI":"10.1038\/srep11783","article-title":"Dietary protein and lifespan across the metamorphic boundary: Protein-restricted larvae develop into short-lived adults","volume":"5","author":"Runagall-McNaull","year":"2015","journal-title":"Scientific Reports"},{"issue":"4","key":"2026101500270531600_bib76","doi-asserted-by":"publisher","first-page":"e71070","DOI":"10.1002\/ece3.71070","article-title":"Evolutionary history with chronic malnutrition enhances pathogen susceptibility at older ages","volume":"15","author":"Sarkar","year":"2025","journal-title":"Ecology and Evolution"},{"key":"2026101500270531600_bib77","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1146\/annurev-ento-010715-023924","article-title":"Reproduction-immunity trade-offs in insects","volume":"61","author":"Schwenke","year":"2016","journal-title":"Annual Review of Entomology"},{"issue":"17","key":"2026101500270531600_bib78","doi-asserted-by":"publisher","first-page":"e2412124122","DOI":"10.1073\/pnas.2412124122","article-title":"Pathogen growth and virulence dynamics drive the host evolution against coinfections","volume":"122","author":"Seal","year":"2025","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2026101500270531600_bib79","doi-asserted-by":"publisher","first-page":"e68874","DOI":"10.7554\/eLife.68874","article-title":"Evolution of pathogen tolerance and emerging infections: A missing experimental paradigm","volume":"10","author":"Seal","year":"2021","journal-title":"eLife"},{"issue":"7","key":"2026101500270531600_bib80","doi-asserted-by":"publisher","first-page":"20250136","DOI":"10.1098\/rsbl.2025.0136","article-title":"Exploring the role of microbiota in mediating sexually dimorphic infection outcomes in mealworm beetles","volume":"21","author":"Seal","year":"2025","journal-title":"Biology Letters"},{"issue":"2","key":"2026101500270531600_bib81","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1159\/000449175","article-title":"The rise and fall of protein malnutrition in global health","volume":"69","author":"Semba","year":"2016","journal-title":"Annals of Nutrition and Metabolism"},{"issue":"1987","key":"2026101500270531600_bib82","doi-asserted-by":"publisher","first-page":"20221642","DOI":"10.1098\/rspb.2022.1642","article-title":"Ageing leads to reduced specificity of antimicrobial peptide responses in Drosophila melanogaster","volume":"289","author":"Shit","year":"2022","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"1700","key":"2026101500270531600_bib83","doi-asserted-by":"publisher","first-page":"3649","DOI":"10.1098\/rspb.2010.0937","article-title":"Female and male genetic contributions to post\u2013mating immune defence in female Drosophila melanogaster","volume":"277","author":"Short","year":"2010","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"135","key":"2026101500270531600_bib84","doi-asserted-by":"publisher","first-page":"57676","DOI":"10.3791\/57676","article-title":"Oral bacterial infection and shedding in Drosophila melanogaster","author":"Siva-Jothy","year":"2018","journal-title":"Journal of Visualized Experiments"},{"key":"2026101500270531600_bib85","doi-asserted-by":"publisher","first-page":"274","DOI":"10.1046\/j.1365-3032.1998.233090.x","article-title":"Decreased immune response as a proximate cost of copulation and oviposition in a damselfly","volume":"23","author":"Siva-Jothy","year":"1998","journal-title":"Physiological Entomology"},{"issue":"16","key":"2026101500270531600_bib86","doi-asserted-by":"publisher","first-page":"3350","DOI":"10.1111\/febs.15327","article-title":"Glycolysis\u2014a key player in the inflammatory response","volume":"287","author":"Soto-Heredero","year":"2020","journal-title":"The FEBS Journal"},{"issue":"10","key":"2026101500270531600_bib87","doi-asserted-by":"publisher","first-page":"1753","DOI":"10.3390\/ijms17101753","article-title":"Critical analysis of the melanogenic pathway in insects and higher animals","volume":"17","author":"Sugumaran","year":"2016","journal-title":"International Journal of Molecular Sciences"},{"issue":"1859","key":"2026101500270531600_bib88","doi-asserted-by":"publisher","first-page":"20170727","DOI":"10.1098\/rspb.2017.0727","article-title":"Dissecting the contributions of time and microbe density to variation in immune gene expression","volume":"284","author":"Tate","year":"2017","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"3","key":"2026101500270531600_bib89","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1093\/infdis\/jis527","article-title":"Protein energy malnutrition decreases immunity and increases susceptibility to influenza infection in mice","volume":"207","author":"Taylor","year":"2013","journal-title":"The Journal of Infectious Diseases"},{"key":"2026101500270531600_bib90","volume-title":"A package for survival analysis in 24R","author":"Therneau","year":"2024"},{"issue":"2","key":"2026101500270531600_bib91","doi-asserted-by":"publisher","first-page":"e1006847","DOI":"10.1371\/journal.ppat.1006847","article-title":"Comparative transcriptomics reveals CrebA as a novel regulator of infection tolerance in D. melanogaster","volume":"14","author":"Troha","year":"2018","journal-title":"PLOS Pathogens"},{"issue":"6","key":"2026101500270531600_bib92","doi-asserted-by":"publisher","first-page":"1512","DOI":"10.1016\/j.cell.2016.07.026","article-title":"Opposing effects of fasting metabolism on tissue tolerance in bacterial and viral inflammation","volume":"166","author":"Wang","year":"2016","journal-title":"Cell"},{"key":"2026101500270531600_bib93","doi-asserted-by":"publisher","DOI":"10.55430\/6304TDIHVA01","volume-title":"The Drosophila immunity handbook","author":"Westlake","year":"2024"},{"key":"2026101500270531600_bib94","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-24277-4","volume-title":"ggplot2: Elegant graphics for data analysis","author":"Wickham","year":"2016"},{"key":"2026101500270531600_bib95","doi-asserted-by":"publisher","first-page":"1183659","DOI":"10.3389\/fgene.2023.1183659","article-title":"Lipids as a key element of insect defense systems","volume":"14","author":"Wro\u0144ska","year":"2023","journal-title":"Frontiers in Genetics"},{"issue":"5","key":"2026101500270531600_bib96","doi-asserted-by":"publisher","first-page":"888","DOI":"10.1016\/j.cmet.2021.01.017","article-title":"Very-low-protein diets lead to reduced food intake and weight loss, linked to inhibition of hypothalamic mTOR signaling, in mice","volume":"33","author":"Wu","year":"2021","journal-title":"Cell Metabolism"},{"issue":"1","key":"2026101500270531600_bib97","doi-asserted-by":"publisher","first-page":"e1000267","DOI":"10.1371\/journal.ppat.1000267","article-title":"The sicker sex","volume":"5","author":"Zuk","year":"2009","journal-title":"PLoS Pathogens"},{"issue":"10","key":"2026101500270531600_bib98","doi-asserted-by":"publisher","first-page":"1009","DOI":"10.1016\/S0020-7519(96)80001-4","article-title":"Sex differences in parasite infections: Patterns and processes","volume":"26","author":"Zuk","year":"1996","journal-title":"International Journal for Parasitology"}],"container-title":["Evolution"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/evolut\/advance-article-pdf\/doi\/10.1093\/evolut\/qpag175\/71396120\/qpag175.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/evolut\/advance-article-pdf\/doi\/10.1093\/evolut\/qpag175\/71396120\/qpag175.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T04:27:15Z","timestamp":1792038435000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/evolut\/advance-article\/doi\/10.1093\/evolut\/qpag175\/8835803"}},"subtitle":[],"editor":[{"given":"Ruth","family":"Archer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]},{"given":"Jason","family":"Wolf","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,9,25]]},"references-count":98,"URL":"https:\/\/doi.org\/10.1093\/evolut\/qpag175","relation":{},"ISSN":["0014-3820","1558-5646"],"issn-type":[{"value":"0014-3820","type":"print"},{"value":"1558-5646","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,25]]},"article-number":"qpag175"}}