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Adaptive significance of Non Genetic Inheritance (ANGI) Dr. Benoit PUJOL Host institution: Contents: Context Rationale Objectives Approach Project management EDB Lab - Toulouse, France How selection shapes diversity? Functional Ecology Evolutionary biology Quantitative genetics The ability of natural populations to adapt environmental constraints 1 Context & rationale – Objectives & Approach – Project management PUJOL – ANGI project Implications of living on earth EARTH © Wikimedia Commons / based upon a NASA image 2 Context & rationale – Objectives & Approach – Project management PUJOL – ANGI project It is changing right now © NASA 3 Context & rationale – Objectives & Approach – Project management PUJOL – ANGI project Sea level rises 10-20 cm over 100y Annual rise: 3.3 mm.y-1 btw 1993 – 2009 Likely doubled since 2014 33 to 120cm in 2100 You live in Camargue: Learn free diving! 4 Context & rationale – Objectives & Approach – Project management PUJOL – ANGI project What solutions? Imagine you are adapted to live in the arctic circle Good fitness = reproduction*survival = species maintenance © Wikimedia commons 6 Context & rationale – Objectives & Approach – Project management PUJOL – ANGI project Solution 1: do nothing about it do like humans.. Not much.. Maybe Bruce willis will save the world again © Africa geographic, Michael C. Scholl and Thomas P. Peschak © photo by Gage Skidmore 7 Context & rationale – Objectives & Approach – Project management PUJOL – ANGI project Solution 2: Migrate better catch the last icebreaker going up north! © Wikimedia commons 8 Context & rationale – Objectives & Approach – Project management PUJOL – ANGI project Run you fools… On average, species distribution go up 11m per decade © Wikimedia commons / Oregon’s Mt. Hood territory Metanalysis by Chen et al. 2011 They also go 16.9km north per decade © Wikimedia commons / Pearson Scott Foresman 9 Context & rationale – Objectives & Approach – Project management PUJOL – ANGI project Solution 3: change, adapt, evolve © Wikimedia commons 10 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project How do you adapt? Survival of the fittest Natural selection Charles Darwin Alfred Wallace 11 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project Genetic variation = response to selection Breeder’s equation R=h²S Assumptions = infinitesimal model, linear additive covariance relation (h²=VA/VP) The example of human height (h²=0.83) … 12 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project Itininéraire d’un enfant gaté Pas à pas, les atouts biologie intégrative et expérimentation végétale sortent de la manche … ANR JCJC CAPA Adaptation aux changements environnementaux Projet « new frontiers » Plasticité phénotypique IDEX CAPS Migration en populations naturelles Our ability to predict adaptive evolution is limited Selective breeding Triticum sp. Zea mays Natural selection Adaptation to environmental change Evidence for genetic response Drosophila Brassica rapa Bos taurus Gallus gallus Highly predictable response when environmental variation is excluded Fagus sylvatica Wyeomyia smithii In natural populations, the predicted genetic response to selection is often missing 13 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project In the wild: adaptive plasticity Parus major Perisoreus infaustus Arabidopsis Beta vulgaris Tamiasciurus hudsonicus Chroicocephalus scopulinus environmentally induced phenotypic response that allow species to acclimatise When adaptive phenotypic changes are observed: 4 1/3 only was caused by genetic change Gienapp et al 2007 (Mol Ecol) Raises questions about the mechanisms shaping observed patterns of phenotypic plasticity Ficedula albicollis 14 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project That’s where our project kicks in.. Adaptive significance of Non Genetic Inheritance Laboratoire Toulouse, France This project has received funding from the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement No [681484] ANGI) L’hérédité génétique qu’est ce que c’est? Pesticides, polluants environnementaux, perturbateurs endocriniens Atteint directement la lignée germinale de l’embryon (méthylation de l’ADN) Chute de la fertilité mâle (moins de spermatocytes) Sur au moins 4 générations sans amélioration… Tant que ça reste une histoire de souris … Implication possible pour la santé ... Dommageable pour les protéines qui lient l’ADN et l’empêche de se casser Cas particulier, on ne connait pas l’impact sur les populations? Concept à implications sociétales L’hérédité, qu’est ce que c’est? La réponse d’« expert » : « J’inclinerais, pour ma part, à penser qu’on naît pédophile, et c’est d’ailleurs un problème que nous ne sachions soigner cette pathologie. Il y a mille deux cents ou mille trois cents jeunes qui se suicident en France chaque année, ce n’est pas parce que leurs parents s’en sont mal occupés ! Mais parce que, génétiquement, ils avaient une fragilité, une douleur préalable. » L’importance de sortir du dogme du tout génétique se passe de commentaires.. Ecological significance Transmission of the environmentally induced phenotypic response NGI influences ecological dynamics (Nat Rev Genet 2011) Example: ecological niche transmission Castor sp. Lodge (pond and protection) Odling-Smee et al. 2003 (Book) Campanulastrum americanum Galloway et al. 2007 (science) shade effects on the phenotype Concern: studies confounded by genetic [DNA] heterogeneity (temporal and geographic level) 15 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project Methodological challenge Double pedigree method: Separate genetic and non genetic covariance with phenotypic resemblance (2013 PLoS ONE) Phenotypic variation Inherited variation Genetic variation (VG) Non-transmitted variation Non genetically transmitted variation (VNG) 17 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project Evolutionary implications of NGI Quantitative genetics Classical theory NGI Current view NGI trait variation Genetic trait variation Selection pressure Proportional response to selection Similar Selection pressure Proportional response to selection Otto et al. 1995 (Stanford working paper series) Day and Bonduriansky 2011 (Am Nat) Townley and Ezard 2013 (J evolution Biol) 18 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project Evolutionary implications of NGI Gap in the theory Absence of empirical validation (NGI-Fitness relationship) Adaptive significance is still debated Current view NGI trait variation Fitness link Selection pressure Proportional response to selection 19 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project Evolutionary implications of NGI Critical assumptions must be included Modification of predictive models Feed back concept model validation Current view NGI trait variation Non mendelian Non vertical Fitness link Selection pressure Proportional response to selection Only partly conserved 20 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project Aim of ANGI project: assess the role of NGI in adaptive evolution Three objectives: 1. Empirical test in the wild expectation: non genetically inherited variation in fitness WP1 WP2 (adaptive evolution) WP3 2. Experimental selection: Generate epigenetic trait variation and test for its response to selection (and stability) WP4 3. Revisit current theory Contribute to develop an explicit quantitative non-genetic theory of natural selection WP5 21 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project The model system Antirrhinum majus (snapdragon) Model species (abandoned salt marsh – Nature Park) Fragmented populations (n=6) (7y, n~7000, location, environment, phenotype) Multiple traits with ecological relevance (morphology, reproduction, predation) Data for a large number of replicated microhabitat patches 22 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project stem elongation in response to shade School book example 23 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project Project management Building on Previous work Funding Juliette, Sara, Aurore Beyond classical quantitative genetics Mathieu Isabel, Caroline, Mathilde, Thank you for your attention 27 Context & rationale – Objectives & Approach – Project managementPUJOL – ANGI project