Flatt lab publications

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Publication list in pdf format



Promislow DEL*, Flatt T*, & Bonduriansky R*. 2022. The Biology of Aging in Insects: From Drosophila to Non-Model Species. [*equal contribution; co-corresponding authors]. Annual Review of Entomology (planned for Volume 67, 2022), in press.


Betancourt N, Rajpurohit S, Durmaz E, Fabian DK, Kapun M, Flatt T*, & Schmidt P*.  2021. Allelic polymorphism at foxo contributes to local adaptation in Drosophila melanogaster [*co-corresponding authors]. Molecular Ecology, in press. 


Wallace M, Coffman KA, Gilbert C, Ravindran S, Albery GF, Argyridou E, Bellosta P, Betancourt A, Colinet H, Eric K, Glaser-Schmitt A, Grath S, Jelic M, Kankare M, Kozeretska I, Loeschcke V, Montchamp-Moreau C, Ometto L, Onder BS, Orengo DJ, Parsch J, Pascual M, Patenkovic A, Puerma E, Ritchie MG, Rota-Stabelli O, Schou MF, Serga SV, Stamenkovic-Radak M, Tanaskovic M, Veselinovic MS, Vieira J, Vieira CP, Kapun M, Flatt T, González J, Staubach F, & Obbard DJ. 2021. The discovery, distribution and diversity of DNA viruses associated with Drosophila melanogaster in Europe. Virus Evolution, in press. 


Korb J, Meusemann K, Aumer D, Bernadou A, Elsner D, Feldmeyer B, Foitzik S, Heinze J, Libbrecht R, Lin S, Majoe M, Monroy Kuhn JM, Nehring V, Negroni M, Paxton R, Séguret A, Stoldt M, & Flatt T (2021). Comparative Transcriptomic Analysis of the Mechanisms Underpinning Ageing and Fecundity in Social Insects. Philosophical Transactions of the Royal Society of London B, 376 (1823):20190728. pdf [Also see coverage in Science, 25 March, 2021, https://doi.org/10.1126/science.371.6536.1302].


Pen I, & Flatt T (2021). Asymmetry, Division of Labour and the Evolution of Ageing in Multicellular Organisms. Philosophical Transactions of the Royal Society London B, 376 (1823):20190729. pdf [Also see coverage in Science, 25 March, 2021, https://doi.org/10.1126/science.371.6536.1302].



Durmaz E, Kerdaffrec E, Katsianis G, Kapun M*, & Flatt T* (2020). How Selection Acts on Chromosomal Inversions. [*co-corresponding]. eLS (Encyclopedia of Life Sciences) 1:307-315, 2020. pdf


Kapun M*, Barrón MG, Staubach F, Vieira J, Obbard DJ, Wiberg RAW, Goubert C, Rota-Stabelli O, Kankare M, Haudry A, Waidele L, Kozeretska I, Pasyukova EG, Loeschcke V, Pascual M, Vieira CP, Serga S, Montchamp-Moreau C, Abbott J, Gibert P, Porcelli D, Posnien N, Grath S, Sucena E, Bergland AO, Garcia Guerreiro MP, Onder BS, Argyridou E, Guio L, Schou MF, Deplancke B, Vieira C, Ritchie MG, Zwaan BJ, Tauber E, Orengo DJ, Puerma E, Aguadé M, Schmidt PS, Parsch J, Betancourt AJ, Flatt T*, & González J* (2020). Genomic analysis of European Drosophila melanogaster populations reveals longitudinal structure, continent-wide selection, and unknown DNA viruses. [*co-corresponding authors]. Molecular Biology and Evolution, 37:2661-2678. pdf


Waldvogel A-M, Feldmeyer B, Exposito-Alonso M, Rellstab C, Kofler R, Mock T, Schmid K, Schmitt I, Bataillon T, Savolainen O, Bergland AO, Flatt T, Guillaume F, & Pfenninger M (2020). Evolutionary genomics can improve prediction of species’ responses to climate change. Evolution Letters 4:4-18. pdf


Flatt T (2020). Life-History Evolution and the Genetics of Fitness Components in Drosophila melanogaster. Genetics 214:3-48. pdf



Flatt T (2019). How flies turn food into progeny. eLife 2019; e51289. pdf


Hoedjes K, van den Heuvel J, Kapun M, Keller L, Flatt T*, & Zwaan BJ* (2019). Distinct Genomic Signals of Lifespan and Life History Evolution in Response to Postponed Reproduction and Larval Diet in Drosophila. (*co-corresponding authors). Evolution Letters 3:598-609. pdf


Ramaekers A, Weinberger S, Claeys A, Kapun M, Yan J, Wolf R, Flatt T, Buchner E, & Hassan BA (2019). Altering the temporal regulation of one transcription factor drives evolutionary trade-offs between head sensory organs. Developmental Cell 50:780-792. pdf [also see commentary by I Almudi  & AP McGregor (2019). Sensory Organ Size Evolution: A View from Drosophila. Developmental Cell 50:673-674.]


Durmaz E, Rajpurohit S, Betancourt N, Fabian DK, Kapun M, Schmidt P*, & Flatt T* (2019). A clinal polymorphism in the insulin signaling transcription factor foxo contributes to life-history adaptation in Drosophila (*co-corresponding authors). Evolution 73:1774-1792. pdf [also see commentary by O El-Deeb. 2019. Digest: A clinal polymorphism and life-history adaptations in Drosophila. Evolution 73:2026-2027].


May T, van den Heuvel J, Doroszuk A, Hoedjes K, Flatt T, & Zwaan BJ (2019). Adaptation to developmental diet influences the response to selection on age at reproduction in the fruit fly. Journal of Evolutionary Biology, In Press. pdf


Kapun M & Flatt T (2019). The adaptive significance of chromosomal inversion polymorphisms in Drosophila melanogaster. Molecular Ecology 28:1263-1282. pdf


Colchero F, Jones OR, Conde DA, Hodgson D, Zajitschek F, Schmidt BR, Malo AF, Alberts SC, Becker PH, Bouwhuis S, Bronikowski AM, De Vleeschouwer KM, Delahey RJ, Dummermuth S, Fernández-Duque E, Frisenvænge J, Hesselsøe M, Larson S, Lemaître JF, McDonald J, Miller DAW, O’Donnell C, Packer C, Raboy BE, Reading CJ, Wapstra E, Weimerskirch H, While GM, Baudisch A, Flatt T, Coulson T, & Gaillard JM. (2019). The diversity of population responses to environmental change. Ecology Letters 22:342-353. pdf



Fabian DK, Garschall K, Klepsatel P, Santos-Matos G, Sucena E, Kapun M, Lemaitre B, Schlötterer C, Arking R, & Flatt T (2018). Evolution of longevity improves immunity in Drosophila. Evolution Letters 2(6):567-579. pdf


Flatt T & Partridge L (2018). Horizons in the Evolution of Aging. BMC Biology 16(1):93. pdf


Durmaz E, Benson C, Kapun M, Schmidt P & Flatt T (2018). An Inversion Supergene in Drosophila Underpins Latitudinal Clines in Survival Traits. Journal of Evolutionary Biology 31: 1354-1364. pdf


Flatt T & Wagner GP (2018). Canalization and Robustness. Oxford Bibliographies in Evolutionary Biology. pdf


Garschall K & Flatt T (2018). The interplay between immunity and aging in Drosophila. F1000Research, 7(F1000 Faculty Rev):160. pdf


Andreatta G, Kyriacou CP, Flatt T* & Costa R* (2018). Aminergic signaling controls ovarian dormancy in Drosophila. Scientific Reports 8:2030, 1-14. (*co-corresponding authors) pdf



Hoedjes KM, Rodrigues MA & Flatt T (2017). Amino acid modulation of lifespan and reproduction in Drosophila. Current Opinion in Insect Science 23: 118-122. pdf


Kubrak OI, Nylin S, Flatt T, Nässel DR, & Leimar O (2017). Adaptation to fluctuating environments in a selection experiment with Drosophila melanogaster. Ecology & Evolution 7: 3796–3807. pdf


Garschall K, Dellago H, Gáliková M, Schosserer M, Flatt T* & Johannes Grillari J* (2017). Ubiquitous overexpression of the DNA repair factor dPrp19 reduces DNA damage and extends Drosophila life span. npj Aging and Mechanisms of Disease 3(5). (*co-corresponding authors). pdf



Rodrigues MA & Flatt T (2016). Endocrine uncoupling of the trade-off between reproduction and somatic maintenance in eusocial insects. Current Opinion in Insect Science 16:1-8. pdf


Kapun M, Schmidt C, Durmaz E, Schmidt PS & Flatt T (2016). Parallel effects of the inversion In(3R)Payne on body size across the North American and Australian clines in Drosophila melanogaster. Journal of Evolutionary Biology 29:1059-1072. pdf


Kapun M, Fabian DK, Goudet J & Flatt T (2016). Genomic evidence for adaptive inversion clines in Drosophila melanogaster. Molecular Biology and Evolution 33:1317-1336. pdf


Flatt T (2016). Genomics of clinal variation in Drosophila: disentangling the interactions of selection and demography. Molecular Ecology 25:1023-1026. pdf


Fuellen G, Schofield PN, Flatt T, Schulz R-J, Boege F, Kraft K, Rimbach G, Ibrahim S, Tietz A, Schmidt C, Köhling R & Simm A (2016). Living long and well: prospects for a personalized approach to the medicine of ageing. Gerontology 62: 409-416. pdf



Flatt T (2015). Paying the costs of reproduction. eLife 2015;4:e09556. pdf


Fabian D K, Lack JB, Mathur V, Schlötterer C, Schmidt PS, Pool JE & Flatt T (2015). Spatially varying selection shapes life history clines among populations of Drosophila melanogaster from sub-Saharan Africa. Journal of Evolutionary Biology 28:826-840. pdf



Flatt T (2014). Plasticity of lifespan – A reaction norm perspective. Proceedings of the Nutrition Society 73:532-542. pdf


Klepsatel P, Galikova M, Huber CD &  Flatt T (2014). Similarities and differences in altitudinal versus latitudinal variation for morphological traits in Drosophila melanogaster. Evolution 68:1385- 1398. pdf


Kapun M, van Schalwyk H, McAllister B, Flatt T & Schlötterer C (2014). Inference of chromosomal inversion dynamics from Pool-Seq data in natural and laboratory populations of D. melanogaster. Molecular Ecology 23:1813-1827 [also see commentary by A Navarro & R Faria (2014). Pool and conquer: new tricks for (c)old problems. Molecular Ecology 23:1653-1655]. pdf



Klepsatel P, Galikova M, De Maio N, Huber C, Schlötterer C & Flatt T (2013). Variation in thermal performance and reaction norms among populations of Drosophila melanogaster. Evolution 67:3573-3587. pdf


Flatt T, Amdam GV, Kirkwood TBL & Omholt SW (2013). Life-history evolution and the polyphenic regulation of somatic maintenance and survival. Quarterly Review of Biology 88:185- 218. pdf


Alcedo J, Flatt T & Pasyukova E (2013). The role of the nervous system in aging and longevity. Frontiers in Genetics 4:124 (doi: 10.3389/fgene.2013.00124). pdf


Rus F, Flatt T, Tong M, Kleino A, Aggarwal K, Okuda K, Yates E, Tatar M & Silverman N (2013). Ecdysone triggered PGRP-LC expression controls Drosophila innate immunity. EMBO Journal 32:1626-1638. pdf


Klepsatel P, Galikova M, De Maio N, Ricci S, Schlötterer C & Flatt T (2013). Reproductive and post-reproductive life history of wild-caught Drosophila melanogaster. Journal of Evolutionary Biology 26:1508-1520. pdf


Alcedo J*, Flatt T* & Pasyukova E* (2013). Neuronal inputs and outputs of aging and longevity. Frontiers in Genetics 4:71 (doi: 10.3389/fgene.2013.00071). (*co-corresponding authors) pdf


Hansen M*, Flatt T* & Aguilaniu H* (2013). Reproduction, fat metabolism, and life span: what is the connection? Cell Metabolism 17:10-19. (*co-corresponding authors) pdf



Fabian DK, Kapun M, Nolte V, Kofler R, Schmidt PS, Schlötterer C & Flatt T (2012). Genome-wide patterns of latitudinal differentiation among populations of Drosophila melanogaster from North America. Molecular Ecology 21:4748–4769. pdf


Flatt T (2012). A new definition of aging? Frontiers in Genetics 3:148. doi:10.3389/fgene.2012.00148. [Commentary on  Rose et al. 2012, see below]. pdf


Rose MR, Flatt T, Graves JL, Greer L, Martinez DE, Matos MM, Mueller LD, Shmookler Reis RJ & Shahrestani P (2012). What is aging? Frontiers in Genetics 3:134.doi: 10.3389/ fgene.2012.00134. pdf


Orozco-terWengel P, Kapun M, Nolte V, Kofler R, Flatt T & Schlötterer C (2012). Adaptation of Drosophila to a novel laboratory environment reveals temporally heterogeneous trajectories of selected alleles. Molecular Ecology 21:4931-4941 [featured on Cover; also see commentary by MK Burke & AD Long (2012). What paths do advantageous alleles take during short-term evolutionary change? Molecular Ecology 21:4913-4916]. pdf


Fabian DK & Flatt T (2012). Life history evolution. Nature Education Knowledge 3(10):24. pdf



Fabian DK & Flatt T (2011). The evolution of aging. Nature Education Knowledge 3(10):9. pdf


Flatt T, Heyland, A & Stearns SC. 2011. What Mechanistic Insights Can or Cannot Contribute to Life History Evolution - An Exchange Between Stearns, Heyland, and Flatt. Chapter 28, pp. 375- 379,  in Flatt T Heyland A (Eds.): Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK. pdf


Flatt T & Heyland A (2011). Life History Integration and Trade-Offs. Part 6, pp. 267-269, n Flatt T Heyland A (Eds.): Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK.


Flatt T & Heyland A (2011). Life History Plasticity. Part 5, pp. 219-220, in Flatt T Heyland A (Eds.): Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK.


Flatt T & Heyland A (2011). Lifespan, Aging and Somatic Maintenance. Part 4, pp. 169-170, in Flatt T Heyland A (Eds.): Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK.


Flatt T & Heyland A (2011). Reproduction. Part 3, pp. 99-100, in Flatt T Heyland A (Eds.): Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK.


Flatt T & Heyland A (2011). Growth, Development, and Maturation. Part 2, pp. 27-28,  in Flatt T Heyland A (Eds.): Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK.


Braendle C, Heyland A & Flatt T (2011). Integrating Mechanistic and Evolutionary Analysis of Life History Variation. Chapter 1, pp. 3-10, in Flatt T Heyland A (Eds.): Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK. pdf


Flatt T & Heyland A (2011). Integrating Mechanisms into Life History Evolution. Part 1, pp. 1-2, in Flatt T Heyland A (Eds.): Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK.


Flatt T & Heyland A (2011), editors. Mechanisms of Life History Evolution. The Genetics and Physiology of Life History Traits and Trade-Offs. Oxford University Press, Oxford, UK. 478 pages, 75 illustrations. ISBN 978-0-19-956877-2. (Book reviews in: Times Higher Education, The Quarterly Review of Biology, American Journal of Human Biology, Journal of Genetics - see book reviews).


Klepsatel P & Flatt T (2011). The genomic and physiological basis of life history variation in a butterfly metapopulation. Molecular Ecology 20:1795-1798. pdf


Flatt T (2011). Survival costs of reproduction in Drosophila. Experimental Gerontology 46:369-375. pdf


Galikova M, Klepsatel P, Senti G & Flatt T (2011). Steroid hormone regulation of C. elegans and Drosophila aging and life history. Experimental Gerontology 46:141-147. pdf



Bronikowski AM & Flatt T (2010). Aging and its demographic measurement. Nature Education Knowledge 3(10):3 pdf


Kapun M, Nolte V, Flatt T & Schlötterer C (2010). Natural host range and host specificity of the Drosophila C virus. PLoS One 5(8):e12421. pdf


Galikova M & Flatt T (2010). Dietary restriction and other lifespan extending pathways converge at the activation of the downstream effector takeout. Aging 2:387-390. pdf


Vogl C, Flatt T, Fuhrmann B, Hofmann E, Wallner B, Kovarik P, Strobl B & Müller M (2010). Transcriptome analysis reveals a major impact of tyrosine kinase 2 (Tyk2) on the expression of interferon responsive and metabolic genes. BMC Genomics 25;11(1):199. pdf



Flatt T (2009). Ageing: diet and longevity in the balance. Nature 462:989-990. pdf


Flatt T & Schmidt PS (2009). Integrating evolutionary and molecular genetics of aging. Biochimica et Biophysica Acta 1790:951-962. pdf


Bauer JH, Morris SNS, Chang C, Flatt T, Wood JG & Helfand SL (2009). dSir2 and Dmp53 interact to mediate aspects of CR-dependent life span extension in D. melanogaster. Aging 1:38-48. [Commentary by L.A. Donehower. 2009. Longevity regulation in flies: A role for p53. Aging 1:6-8.]. pdf



Flatt T, Heyland A, Rus F, Porpiglia E, Sherlock C, Yamamoto R, Garbuzov A, Palli SR, Tatar M & Silverman N (2008). Hormonal regulation of the humoral innate immune response in Drosophila melanogaster. Journal of Experimental Biology 211: 2712-2724. [Image Featured in JEB Calendar 2009]. pdf


Flatt T, Min K-J, D’Alterio C, Villa-Cuesta E, Cumbers J, Lehmann R, Jones DL & Tatar M (2008). Drosophila germ-line modulation of insulin signaling and lifespan. Proceedings of the National Academy of Sciences USA 105:6368-6373. pdf


Crook TC. Flatt T & Smiseth PT (2008). Hormonal modulation of larval begging and growth in the burying beetle Nicrophorus vespilloides. Animal Behaviour 75:71-77. pdf



Flatt T & Promislow DEL (2007). Physiology: still pondering an age-old question. Science 318:1255-1256. pdf


Flatt T & Kawecki TJ (2007). Juvenile hormone as a regulator of the trade-off between reproduction and life span in Drosophila melanogaster. Evolution 61:1980-1991. pdf


Shingleton A, Frankino WA, Flatt T, Nijhout HF & Emlen D (2007). Size and shape: the developmental regulation of static allometry in insects. BioEssays 29:536-548. pdf


Min K-J, Flatt T, Kulaots I & Tatar M (2007). Counting calories in Drosophila dietary restriction. Experimental Gerontology 42:247-251. pdf



Flatt T, Moroz LL, Tatar M & Heyland A (2006). Comparing thyroid and insect hormone signaling. Integrative & Comparative Biology 46:777-794. pdf


Bishop C, Erezyilmaz DF, Flatt T, Georgiou CD, Hadfield M, Heyland A, Hodin J, Jacobs M, Maslakova SA, Pires A, Reitzel A, Santagata S, Tanaka K & Youson JH (2006). What is metamorphosis? Integrative & Comparative Biology 46:655-661. pdf


Braendle C & Flatt T (2006). A role for genetic accommodation in evolution? BioEssays 28:868- 873. pdf


Killingback T, Bieri J & Flatt T (2006). Evolution in group-structured populations can resolve the tragedy of the commons. Proceedings of the Royal Society of London B 273:1477-1481. pdf


Tu MP, Flatt T & Tatar M (2006). Juvenile and steroid hormones in Drosophila melanogaster longevity. Chapter 16 (pp. 415-448) in Masoro E J & Austad SN (Eds.), Handbook of the Biology of Aging, 6th Edition, Academic Press (Elsevier), San Diego. pdf



Flatt T, Tu M-P & Tatar M (2005). Hormonal pleiotropy and the juvenile hormone regulation of Drosophila development and life history. BioEssays 27:999-1010. [Featured on Cover Image]. pdf


Flatt T (2005). The evolutionary genetics of canalization. Quarterly Review of Biology 80:287-316. pdf


Altweg, R, Dummermuth S, Anholt BR & Flatt T (2005). Winter weather affects asp viper (Vipera aspis aspis) population dynamics through susceptible juveniles. Oikos 110:55-66. pdf



Flatt T & Kawecki TJ (2004). Pleiotropic effects of Methoprene-tolerant (Met), a gene involved in juvenile hormone metabolism, on life history traits in Drosophila melanogaster. Genetica 122:141- 160. pdf


Flatt T & Scheuring I (2004). Stabilizing factors interact in promoting host-parasite coexistence. Journal of Theoretical Biology 228:241-249. pdf


Flatt T (2004). Assessing natural variation in genes affecting Drosophila lifespan. Mechanisms of Ageing and Development 125:155-159. pdf



Rauch G, Simon J-C, Chaubet B, Haack L, Flatt T & Weisser WW (2002). The influence of ant attendance on aphid feeding behaviour investigated with the electrical penetration graph technique. Entomologia Experimentalis et Applicata 102:13-20. pdf



Flatt T, Shine R, Borges-Landaez PA & Downes SJ (2001). Phenotypic variation in an oviparous montane lizard: effects of thermal and hydric incubation environments. Biological Journal of the Linnean Society 74:339-350. pdf


Flatt T, Maire N & Doebeli M. 2001. A bit of sex stabilizes host-parasite dynamics. Journal of Theoretical Biology 212:345-354. pdf


2000 and before

Flatt T & Weisser WW (2000). The effects of mutualistic ants on aphid life-history traits. Ecology 81:3522-3529. pdf


Flatt T, Dummermuth S & Anholt BR (1997). Mark-recapture estimates of survival in populations of the asp viper, Vipera aspis aspis. Journal of Herpetology 31:558-564. pdf