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Adaptive genetic variation and plant adaptation to environmental heterogeneity (AVE)

Titel Englisch Adaptive genetic variation and plant adaptation to environmental heterogeneity (AVE)
Gesuchsteller/in Widmer Alexander
Nummer 127155
Förderungsinstrument Sinergia
Forschungseinrichtung Institut für Integrative Biologie Departement Umweltwissenschaften ETHZ
Hochschule ETH Zürich - ETHZ
Hauptdisziplin Botanik
Beginn/Ende 01.04.2010 - 31.03.2014
Bewilligter Betrag 900'000.00
Alle Daten anzeigen

Alle Disziplinen (3)

Disziplin
Botanik
Umweltforschung
Oekologie

Keywords (5)

adaptive genetic variation; adaptation; natural selection; alpine flora; environmental heterogeneity

Lay Summary (Englisch)

Lead
Lay summary

The question how organisms become adapted to their environment is of central interest in biology. The present project investigates the genetic basis of adaptation to harsh and changing environmental conditions in alpine plant species using a population genomic approach.Sessile organisms, such as plants, have to be able to adapt to their environmental conditions, both biotic and abiotic, in order to survive. In some landscapes, and especially in the Alps, environmental conditions can change over small geographic scales. Here, sites with contrasting environmental conditions, brought about for example by differences in exposure, snow cover or bedrock, are often located in close physical proximity and within the reach of contemporary gene-flow. In the face of such environmental heterogeneity, genetic variation is essential for the adaptation of plants to their environment.Natural selection acts on phenotypic variation, much of which presumably has a genetic basis in natural plant populations. Understanding the nature of the genetic variation underlying this phenotypic variation is of long-standing and general interest in biology. Most genetic variation in natural plant populations is presumably neutral, and difficult to distinguish from genetic variation underlying differences in adaptive traits. One approach to identify such adaptive genetic variation is to establish an association between specific environmental conditions and genetic variation using a landscape genetic approach.The present study therefore addresses the following main questions:1. What is the genetic basis of adaptation to environmental heterogeneity ?2. Do the same genes contribute to adaptation in related species ?3. What is the adaptive value of candidate genes ?With respect to these questions we hypothesize that1. major genes contribute substantially to adaptation; 2. the same genes (or gene families) are involved in adaptation to the same enviornmental factor in different species; 3. some of the identified candidate genes cause above-average fitness in their native environment, but below- average fitness in other environments (i.e. display genetic trade-offs).Results from this study will provide new insights into the genetic basis of adaptation and the nature of adaptive genetic variation. This knowledge not only contributes to our understanding of natural variation and adaptation per se, but helps us to predict the effects of climate change on alpine plants and provides a scientifically informed foundation for the conservation of alpine plant biodiversity.

Direktlink auf Lay Summary Letzte Aktualisierung: 21.02.2013

Verantw. Gesuchsteller/in und weitere Gesuchstellende

Mitarbeitende

Publikationen

Publikation
Sequencing of the genus Arabidopsis identifies a complex history of nonbifurcating speciation and abundant trans-specific polymorphism.
Novikova Polina Yu, Hohmann Nora, Nizhynska Viktoria, Tsuchimatsu Takashi, Ali Jamshaid, Muir Graham, Guggisberg Alessia, Paape Tim, Schmid Karl, Fedorenko Olga M, Holm Svante, Säll Torbjörn, Schlötterer Christian, Marhold Karol, Widmer Alex, Sese Jun, Shimizu Kentaro K, Weigel Detlef, Krämer Ute, Koch Marcus A, Nordborg Magnus (2016), Sequencing of the genus Arabidopsis identifies a complex history of nonbifurcating speciation and abundant trans-specific polymorphism., in Nature genetics, 48(9), 1077-82.
A practical guide to environmental association analysis in landscape genomics
Rellstab Christian, Gugerli Felix, Eckert Andrew J., Hancock Angela M., Holderegger Rolf (2015), A practical guide to environmental association analysis in landscape genomics, in Molecular Ecology, 24(17), 4348-4370.
Efficient detection of novel nuclear markers for Brassicaceae by transcriptome sequencing
Stockenhuber Reinhard, Zoller Stefan, Shimizu-Inatsugi Rie, Gugerli Felix, Shimizu Kentaro K., Widmer Alex, Fischer Martin C. (2015), Efficient detection of novel nuclear markers for Brassicaceae by transcriptome sequencing, in PLOS One, 10(e0128181), 1-19.
Evolution of the Selfing Syndrome in Arabis alpina (Brassicaceae).
Tedder Andrew, Carleial Samuel, Gołębiewska Martyna, Kappel Christian, Shimizu Kentaro K, Stift Marc (2015), Evolution of the Selfing Syndrome in Arabis alpina (Brassicaceae)., in PloS one, 10(6), 0126618-0126618.
Female sterility associated with increased clonal propagation suggests a unique combination of androdioecy and asexual reproduction in populations of Cardamine amara (Brassicaceae)
Tedder A., Helling M., Pannell J. R., Shimizu-Inatsugi R., Kawagoe T., van Campen J., Sese J., Shimizu K. K. (2015), Female sterility associated with increased clonal propagation suggests a unique combination of androdioecy and asexual reproduction in populations of Cardamine amara (Brassicaceae), in Annals of Botany, 115(5), 763-776.
Genome-wide quantification of homeolog expression ratio revealed nonstochastic gene regulation in synthetic allopolyploid Arabidopsis.
Akama Satoru, Shimizu-Inatsugi Rie, Shimizu Kentaro K, Sese Jun (2014), Genome-wide quantification of homeolog expression ratio revealed nonstochastic gene regulation in synthetic allopolyploid Arabidopsis., in Nucleic acids research, 1.
Genomics and the origin of species
Seehausen Ole, Butlin Roger K., Keller Irene, Wagner Catherine E., Boughman Janette W., Hohenlohe Paul A., Peichel Catherine L., Saetre Glenn-Peter, Bank Claudia, Braennstroem Ake, Brelsford Alan, Clarkson Chris S., Eroukhmanoff Fabrice, Feder Jeffrey L., Fischer Martin C., Foote Andrew D., Franchini Paolo, Jiggins Chris D., Jones Felicity C., Lindholm Anna K., Lucek Kay, Maan Martine E., Marques David A., Martin Simon H., Matthews Blake (2014), Genomics and the origin of species, in NATURE REVIEWS GENETICS, 15(3), 176-192.
Validation of outlier loci through replication in independent data sets: a test on Arabis alpina
Buehler Dominique, Brodbeck Sabine, Holderegger Rolf, Schnyder Elvira, Gugerli Felix (2014), Validation of outlier loci through replication in independent data sets: a test on Arabis alpina, in Ecology and Evolution, 4(22), 4296-4306.
A road map for molecular ecology
Andrew Rose L., Bernatchez Louis, Bonin Aurélie, Buerkle C. Alex, Carstens Bryan C., Emerson Brent C., Garant Dany, Giraud Tatiana, Kane Nolan C., Rogers Sean M., Slate Jon, Smith Harry, Sork Victoria L., Stone Graham N., Vines Timothy H., Waits Lisette, Widmer Alex, Rieseberg Loren H. (2013), A road map for molecular ecology, in Molecular Ecology, 22(10), 2605-2626.
Challenges in the studies on flowering time: interfaces between phenological researches and the molecular network of flowering genes
Kobayashi Masaki J., Shimizu Kentaro K. (2013), Challenges in the studies on flowering time: interfaces between phenological researches and the molecular network of flowering genes, in Ecological Research, 28, 161.
Pollen, Pollen tube, Selfing, Pistil, Ovule, Endosperm, Embryo sac, Sporangium, Stamen
Shimizu Kentaro K. (2013), Pollen, Pollen tube, Selfing, Pistil, Ovule, Endosperm, Embryo sac, Sporangium, Stamen, in Iwasa Yoh, Kuratani Shigeru, Saitou Naruya, Tsukaya Hirokazu (ed.), Iwanami, Tokyo, 1.
Population genomic footprints of selection and associations with climate in natural populations of Arabidopsis halleri from the Alps
Fischer Martin C., Rellstab Christian, Tedder Andrew, Zoller Stefan, Gugerli Felix, Shimizu Kentaro K., Widmer Alex (2013), Population genomic footprints of selection and associations with climate in natural populations of Arabidopsis halleri from the Alps, in Molecular Ecology, 22(22), 5594-5607.
Ten years of landscape genetics
Manel Stéphanie, Holderegger Rolf (2013), Ten years of landscape genetics, in Trends in Ecology & Evolution, 28, 614-621.
The more the merrier: recent hybridization and polyploidy in cardamine.
Mandáková Terezie, Kovarík Ales, Zozomová-Lihová Judita, Shimizu-Inatsugi Rie, Shimizu Kentaro K, Mummenhoff Klaus, Marhold Karol, Lysak Martin A (2013), The more the merrier: recent hybridization and polyploidy in cardamine., in The Plant cell, 25(9), 3280-95.
Validation of SNP allele frequencies determined by pooled next-generation sequencing in natural populations of a non-model plant species
Rellstab Christian, Zoller Stefan, Tedder Andrew, Gugerli Felix, Fischer Martin C. (2013), Validation of SNP allele frequencies determined by pooled next-generation sequencing in natural populations of a non-model plant species, in PLoS One, 8, e80422.
Adaptive landscape genetics: pitfalls and benefits
Parisod Christian, Holderegger Rolf (2012), Adaptive landscape genetics: pitfalls and benefits, in Molecular Ecology, 21, 3644-3646.
Broad-scale adaptive genetic variation in alpine plants is driven by temperature and precipitation
Manel S., Gugerli F., Thuiller W., Alvarez N., Legendre P., Holderegger F., Gielly L., Taberlet P., IntraBioDiv Consortium (2012), Broad-scale adaptive genetic variation in alpine plants is driven by temperature and precipitation, in Molecular Ecology, 21(15), 3729-3738.
Chapter 12 Overview: the evolution of genes responsible for adaptation
Shimizu Kentaro K. (2012), Chapter 12 Overview: the evolution of genes responsible for adaptation, in Morinaga Shinichi, Kudoh Hiroshi (ed.), Kyoritsu, Tokyo, 189-196.
Chapter 16: Genome duplication: speciation and ecogenomics of polyploid species
Shimizu-Inatsugi Rie, Shimizu Kentaro K. (2012), Chapter 16: Genome duplication: speciation and ecogenomics of polyploid species, in Morinaga Shinichi, Kudoh Hiroshi (ed.), Kyoritsu, Tokyo, 246-262.
Forecasting changes in population genetic structure of Alpine plants in response to global warming
Jay Flora, Manel Stéphanie, Alvarez Nadir, Durand Eric Y., Thuiller Wilfried, Holderegger Rolf, Taberlet Pierre, François Olivier (2012), Forecasting changes in population genetic structure of Alpine plants in response to global warming, in Molecular Ecology, 21, 2354-2368.
Recent loss of self-incompatibility by degradation of the male component in allotetraploid Arabidopsis kamchatica
Tsuchimatsu Takashi, Kaiser Pascal, Yew Chow-Lih, Bachelier Julien, Shimizu Kentaro (2012), Recent loss of self-incompatibility by degradation of the male component in allotetraploid Arabidopsis kamchatica, in PLoS Genetics, 8(7), e1002838.
Coexistence of trichome variation in a natural plant population: a combined study using ecological and candidate gene approaches
Kawagoe T., Shimizu K.K., Kakutani T., Kudoh H. (2011), Coexistence of trichome variation in a natural plant population: a combined study using ecological and candidate gene approaches, in PLoS One, 6(7), 1-12.
Plant sexual reproduction during climate change: gene function in natura studied by ecological and evolutionary systems biology
Shimizu K.K., Kudoh H., Kobayashi M.J. (2011), Plant sexual reproduction during climate change: gene function in natura studied by ecological and evolutionary systems biology, in Annals of Botany, 108, 777-787.
Landscape genetics of plants
Holderegger R, Buehler D, Gugerli F, Manel S (2010), Landscape genetics of plants, in TRENDS IN PLANT SCIENCE, 15(12), 675-683.
Are adaptive loci transferable across genomes of related species? Outlier and environmental association analyses in Alpine Brassicaceae species
Zulliger Deborah, Schnyder Elvira, Gugerli Felix, Are adaptive loci transferable across genomes of related species? Outlier and environmental association analyses in Alpine Brassicaceae species, in Molecular Ecology, 22(6), 1626-1639.
Genome-wide Comparative Analysis of the GRAS Gene Family in Populus, Arabidopsis and Rice
Liu Xuanyu, Widmer Alex, Genome-wide Comparative Analysis of the GRAS Gene Family in Populus, Arabidopsis and Rice, in Plant Molecular Biology Reporter.
Local adaptation (mostly) remains local: reassessing environmental associations of climate-related candidate SNPs in Arabidopsis halleri
Rellstab Christian, Fischer Martin C., Zoller Stefan, Graf René, Tedder Andrew, Shimizu Kentaro K., Widmer Alex, Holderegger Rolf, Gugerli Felix, Local adaptation (mostly) remains local: reassessing environmental associations of climate-related candidate SNPs in Arabidopsis halleri, in Heredity, online.

Wissenschaftliche Veranstaltungen

Aktiver Beitrag

Titel Art des Beitrags Titel des Artikels oder Beitrages Datum Ort Beteiligte Personen
European Society of Evolutionary Biology Poster Making more out of big data — cost-effective development of novel nuclear markers from transcriptome data (Poster presented by R. Stockenhuber) 10.08.2015 Lausanne, Schweiz Shimizu Kentaro;
Evolution Vortrag im Rahmen einer Tagung Using pooled sequencing and whole-genome environmental association analyses to study local adaptation in three Alpine Brassicaceae species 20.06.2014 Raleigh, Vereinigte Staaten von Amerika Rellstab Christian;
Disease, immunity and ancient DNA Vortrag im Rahmen einer Tagung Evolutionary genomics of mating systems 17.02.2014 Zurich, Schweiz Shimizu Kentaro;
Collaborative visit of delegation from Technical University of Zvolen Einzelvortrag How molecular-genetic tools help understanding ecological processes (contribution of F. Gugerli) 06.02.2014 Birmensdorf, Schweiz Holderegger Rolf;
Symposium of Genetic Diversity Center Vortrag im Rahmen einer Tagung Using pooled next-generation sequencing to determine SNP allele frequencies in natural populations of a non-model plant species (with contribution of F. Gugerli) 30.01.2014 Zürich, Schweiz Holderegger Rolf; Fischer Martin; Tedder Andrew; Rellstab Christian;
Kickoff meeting of the Memorandum of Understanding between Forest Research Institute Malaysia and Faculty of Science, University of Zurich Einzelvortrag Evolutionary and ecological genomics from Arabidopsis to Dipterocarpaceae 20.01.2014 Forest Research Institute Malaysia, Malaysia Shimizu Kentaro;
Mini-symposium on Mating system transitions and their implications for evolutio Vortrag im Rahmen einer Tagung Emergent androdioecy in Swiss Cardamine amara (Brassicaceae) 10.10.2013 Konstanz, Deutschland Tedder Andrew; Shimizu Kentaro;
Mating system transitions and their implications for evolution Vortrag im Rahmen einer Tagung Emergent androdioecy in Swiss Cardamine amara (Brassicaceae) 10.10.2013 Constance, Deutschland Tedder Andrew;
Congress on past, present and future dynamics of alpine and arctic flora under climate change Vortrag im Rahmen einer Tagung Monitoring and forecasting genetic adaptation to climate change in alpine plants (with contributions by S. Zoller and F. Gugerli) 22.09.2013 Bergün, Schweiz Holderegger Rolf; Tedder Andrew; Fischer Martin; Rellstab Christian; Widmer Alexander; Shimizu Kentaro;
Seminar at INRA Centre Bordeaux-Aquitaine, France Einzelvortrag Adaptive genetic variation and adaptation to environmental heterogeneity in Alpine Brassicaceae species (with contribution by F. Gugerli) 04.06.2013 Pierroton, Frankreich Rellstab Christian; Holderegger Rolf;
2nd Arabis Workshop Vortrag im Rahmen einer Tagung Mating system variation and sporophytic self-incompatibility in European Arabis alpina 17.04.2013 Birmensdorf, Schweiz Tedder Andrew;
2nd Arabis Workshop Vortrag im Rahmen einer Tagung Adaptive genetic variation and adaptation to environmental heterogeneity in Alpine Brassicaceae species (with contribution of F. Gugerli) 17.04.2013 Birmensdorf, Schweiz Fischer Martin; Shimizu Kentaro; Widmer Alexander; Holderegger Rolf; Tedder Andrew; Rellstab Christian;
FroSpects Speciation workshop Poster Genome evolution of an Alpine plant in a highly heterogeneous environment 26.03.2013 Kastanienbaum, Schweiz Fischer Martin;
FroSpects Speciation workshop Vortrag im Rahmen einer Tagung Population genomics of adaptation or the complexity of differential adaptation at the genic level 26.03.2013 Kastanienbaum, Switzerland, Schweiz Widmer Alexander; Fischer Martin;
Symposium on Arabidopsis natural accessions and related species, The 54th Annual Meeting of the Japanese Society of Plant Physiologists Vortrag im Rahmen einer Tagung Recurrent patterns in the adaptation and speciation in Arabidopsis relatives 22.03.2013 Okayama, Japan Shimizu Kentaro;
DEG seminar, WSL Einzelvortrag Population genomics of Arabidopsis halleri: patterns of genetic variation and evidence for adaptation to diverse Alpine environments 26.02.2013 Birmensdorf, Switzerland, Schweiz Fischer Martin;
University of Trier Einzelvortrag Next-generation sequencing in non-model organisms - insights from plants 21.02.2013 Trier, Deutschland Widmer Alexander;
Plant and Animal Genome XXI Vortrag im Rahmen einer Tagung A genome-wide perspective on variation and selection in populations of Arabidopsis halleri from the Alps 12.01.2013 San Diego, USA, Vereinigte Staaten von Amerika Fischer Martin;
The 13th Swiss-Japanese conference on biotechnology and bioprocess engineering Vortrag im Rahmen einer Tagung Next-generation sequencing for evolutionary genomics of model and non-model species 11.11.2012 Walzenhausen, Schweiz Shimizu Kentaro;
5th URPP Systems Biology and Functional Genomics Meeting Vortrag im Rahmen einer Tagung Evolutionary and ecological genomics for Arabidopsis relatives and non-model species 14.09.2012 Brunnen, Schweiz Shimizu Kentaro;
Molecular Ecology symposium Einzelvortrag Reproductive isolation, hybridisation and genomic divergence in a dioecious plant group 06.07.2012 Ottawa, Canada, Kanada Widmer Alexander;
Evolution Ottawa Vortrag im Rahmen einer Tagung Population re-sequencing reveals variation and selection in natural 06.07.2012 Ottawa, Canada, Kanada Fischer Martin;
Symposium of Genetic Diversity Center Vortrag im Rahmen einer Tagung Population re-sequencing reveals variation and selection in natural 02.07.2012 Zürich, Switzerland, Schweiz Fischer Martin;
Plant Population Biology of GfÖ Poster Population re-sequencing reveals the signature of habitat-mediated selection in Arabidopsis halleri. 17.05.2012 Zürich, Schweiz Holderegger Rolf;
Seminar Institute of Biology, University of Zürich Einzelvortrag Landscape genomics & local adaptation in alpine plants 16.04.2012 Zürich, Schweiz Rellstab Christian;
Theoretical and empirical advances in evolutionary genomics Poster Population re-sequencing reveals variation and selection in natural populations of Arabidopsis halleri 31.03.2012 Roscoff, France, Frankreich Fischer Martin;
EcoChange final conference Poster Are adaptive loci transferable across genomes of related species? – Outlier and association analyses in five Alpine Brassicaceae species (with contribution by F. Gugerli) 22.03.2012 Zürich, Switzerland, Schweiz Zulliger Deborah; Rellstab Christian;
International Okazaki Biological Conference 8 on Speciation and adaptation Vortrag im Rahmen einer Tagung Toward recurrent patterns in the evolution of breeding systems and polyploid speciation in Arabidopsis relatives 20.03.2012 Okazaki, Japan, Japan Shimizu Kentaro;
The 53th Annual Meeting of the Japanese Society of Plant Physiologists Vortrag im Rahmen einer Tagung Transcriptomic study of polyploid Arabidopsis relatives 17.03.2012 Kyoto, Japan, Japan Shimizu Kentaro;
ESF ConGenOmics workshop Einzelvortrag Developing a genome-wide perspective on genetic variation, adaptation and gene expression in non-model plant species 09.02.2012 Sevilla, Spanien Widmer Alexander;
Adaptive landscape genetics: current insights and future directions Vortrag im Rahmen einer Tagung Population re-sequencing reveals the signature of habitat-mediated selection in Arabidopsis halleri 07.02.2012 Neuchâtel, Switzerland, Schweiz Fischer Martin;
Adaptive Landscape Genetics Vortrag im Rahmen einer Tagung The needle in the haystack: ecologically relevant alleles in non-model plants from the Alps (Keynote by F. Gugerli) 07.02.2012 Neuchatel, Schweiz Holderegger Rolf;
Detection of genomic regions under natural selection in landscape genetics Einzelvortrag Identifying genomic regions showing signals of selection: from outlier and environmental association studies to sequence characterization (contribution of F. Gugerli) 09.12.2011 Lausanne, Schweiz Zulliger Deborah;
European Ecological Federation Poster Identifying adaptive genetic diversity in relation to environmental variation: association genetics in Brassicaceae (with contribution by F. Gugerli) 25.09.2011 Avila, Spain, Spanien Zulliger Deborah;
Institute seminar EAWAG, Kastanienbaum Einzelvortrag Adaptive genetic variation in a landscape context 30.05.2011 Kastanienbaum, Schweiz Holderegger Rolf;
Biology 11 Vortrag im Rahmen einer Tagung Identification and verification of candidate locus for habitat-related adaptation in the alpine plant Arabis alpina (with contribution by F. Gugerli) 03.02.2011 Zürich, Schweiz Holderegger Rolf;
Biology 2011 Poster Identifying adaptive genetic diversity in relation to environmental variation: association genetics in Brassicaceae (with contribution by F. Gugerli) 03.02.2011 Zürich, Switzerland, Schweiz Zulliger Deborah;
Latsis Symposium Vortrag im Rahmen einer Tagung Potential habitat-related adaptation in the alpine plant, Arabis alpina (with contribution by F. Gugerli) 15.11.2010 Zürich, Switzerland, Schweiz Holderegger Rolf;
Evolutionary and Ecological Genomics of Adaptation Poster Multi-species environmental association study identifies loci primarily affected by temperature and precipitation in Alpine plants (contribution by F. Gugerli) 02.09.2010 Fribourg, Switzerland, Schweiz Holderegger Rolf;
Evolutionary and Ecological Genomics of Adaptation Poster Outlier analysis identifies candidate locus for habitat-related adaptation in the alpine plant Arabis alpina (with contribution by F. Gugerli) 02.09.2010 Fribourg, Schweiz Holderegger Rolf;
Forest Ecosystem Genomics and Adaptation Vortrag im Rahmen einer Tagung Identifying adaptive genetic diversity in relation to environmental variation: promising boulevard, but peppered with pitfalls (contribution by F. Gugerli) 09.06.2010 San Lorenzo de El Escorial, Spain, Spanien Holderegger Rolf;


Selber organisiert

Titel Datum Ort
Arabis workshop 17.04.2013 WSL, Birmensdorf, Switzerland, Schweiz
Adaptive landscape genetics: current insights and future directions 07.02.2012 Neuchatel, Switzerland, Schweiz

Verbundene Projekte

Nummer Titel Start Förderungsinstrument
137084 Adaptation and biodiversity in alpine habitats - integrating genomic and ecological perspectives 01.07.2012 ProDoc
158542 PSC Discovery Program for Youth 01.04.2015 Agora
141260 Genomic divergence in plants: exploring the consequences of ecological adaptation for plant divergence and speciation 01.05.2012 Projektförderung (Abt. I-III)
140917 Recurrent patterns in molecular adaptation and speciation: evolutionary genomic analysis using Arabidopsis relatives 01.06.2012 Projektförderung (Abt. I-III)
149741 GENESCALE: Very high-resolution digital elevation models for multi-scale analysis in landscape genomics 01.04.2014 Interdisziplinäre Projekte

Abstract

Natural variation is ubiqitous and forms the target of selection. In sessile organisms, such as plants, strong selection can be imposed by environmental conditions, both biotic and abiotic. These environmental conditions can change over small geographic scales in some landscapes, such as mountain areas and the Alps. Here, sites with contrasting environmental conditions, brought about for example by differences in exposure, snow cover or bedrock, are often located in close physical proximity and within the reach of contemporary gene-flow. In the face of such environmental heterogeneity, natural variation is essential for the adaptation of plants to their environment.The genetic basis of adaptation is one of the central and most hotly debated topics in evolutionary biology. Recent theoretical and empirical work has shed new light on the number and relative effects of genes underlying adaptation. In contrast to earlier views of adaptation being a slow process mediated by large numbers of genes of minor effect, it has become clear that adaptation is often due to a finite number of genes, some of which are of major effect. Empirical studies on adaptation and its underlying adaptive variation in plants have used both forward and reverse genetic approaches, mostly in model organisms, to identify candidate genes for adaptation. The current adaptive value of such candidate genes can in principle be assessed experimentally in transplant experiments. Yet, this ecological work has often been neglected or carried out under highly artificial conditions, mainly because of a lack of collaboration between ecologists and plant molecular biologists. Only a small number of studies, mostly on the model plant species Arabidopsis thaliana, have carefully investigated the adaptive value of candidate genes in different, ecologically realistic environments. These studies have provided new insights into the genetic basis of adaptation, and the role of epistasis, pleiotropy and gene-by-environment interactions in the maintenance of adaptive genetic variation.The presented project builds on the insights gained from such studies to investigate adaptation to environmental heterogeneity from an ecological, landscape genetic and functional genomic perspective. Environmental heterogeneity is pronounced in alpine areas, where conditions often change over small geographic distances as a consequence of differences for example in altitude, exposition, bedrock, snow cover or water availability. Alpine areas are centers of biodiversity in Europe and around the world, and are particularly affected by climate change, yet plant species native to these alpine areas remain poorly investigated with respect to the genetics of adaptation.We here propose to investigate adaptive genetic variation in the model plant species Arabidopsis thaliana and several related species with contrasting mating systems, life histories or altitudinal distributions. We will assess natural genetic variation at 500 loci in multiple populations of six plant species collected in a landscape with high environmental heterogeneity. The 500 loci include 100 candidate genes from A. thaliana and 400 randomly chosen genes that provide approximately homogeneous genome coverage in A. thaliana. This comparative approach is supported by the application of ultra-high throughput sequencing technology, which allows for genome-wide analyses of genetic variation in multiple species.The overarching questions we address are:1.What is the genetic basis of adaptation to environmental heterogeneity ?2.Do the same genes contribute to adaptation in related species ?3.What is the adaptive value of candidate genes ?With respect to these questions we hypothesize that 1.major genes contribute substantially to adaptation2.the same genes (or gene families) are involved in adaptation to the same enviornmental factor in different species3.some of the identified candidate genes cause above-average fitness in their native environment, but below- average fitness in other environments (i.e. display genetic trade-offs).Results from this study will provide new insights into the genetic basis of adaptation and the nature of adaptive genetic variation. This knowledge not only contributes to our understanding of natural variation and adaptation per se, but helps us to predict the effects of climate change on alpine plants and provides a scientifically informed foundation for the conservation of alpine plant biodiversity.
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