Local genetic adaptation to drought: oak species as an example

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1 Local genetic adaptation to drought: oak species as an example Oliver Gailing 1,2, Jennifer Lind 2, Barbara Vornam 1, Reiner Finkeldey 1 1 University of Göttingen, Germany 2 Michigan Technological University, United States 1

2 Adaptive genetic variation in oaks General Introduction Quercus and genomic resources Genetic mapping of drought related traits Species differentiation and local adaptation Outlook 2

3 Adaptive genetic variation in oaks General Introduction Quercus and genomic resources Genetic mapping of drought related traits Species differentiation and local adaptation Outlook 3

4 Introduction climate change Bud phenolgy Temperature increase Longer growing season Changes in the time of bud burst 4 Frost and drought injuries Insect damages Roeckner et al. (2006) Climate Projections for the 21st century. Max Planck Institute for Meteorology.

5 Introduction climate change Stomatal traits, WUE dry summers Drought stress Lower growth Lower biomass 5 Roeckner et al. (2006) Climate Projections for the 21st century. Max Planck Institute for Meteorology.

6 Introduction tree species migration Forest habitats in 2100 Oak/hickory habitat Maple/ beech/birch habitat future species composition 6 Mohan et al. 2009

7 Introduction tree species migration Quercus rubra, Current and predicted distribution 7 A Climate Change Atlas for 80 Forest Tree Species of the Eastern United States [spatial database], Anantha M. Prasad and Louis R. Iverson, USDA Forest Service, Northeastern Research Station, Delaware, Ohio 43015, USA,

8 Adaptive genetic variation in oaks General Introduction Quercus and genomics Adaptive genes and species differentiation Genetic mapping of adaptive traits Outlook 8

9 Quercus as a model Quercus is an excellent model for ecological genetics! 9

10 Quercus as a model Sessile oak (Quercus petraea) Central and Eastern Europe 10

11 Quercus as a model Quercus rubra, Northern Red Oak Barnes and Wagner,

12 12 Adaptive genetic variation in oaks general approach

13 Quercus and genomics EST-SSRs SNPs 13 Jera 1.3; Comparative mapping in Fagaceae Gibson and Muse 2004

14 Quercus and genomics Genomic resources Querus petraea/robur > ESTs 283 EST-SSRs Quercus rubra ESTs Quercus alba ESTs 14 Derory et al. (2006). New Phytologist 170:

15 Quercus and genomics Bin mapping in Quercus robur 4F 4M 5F 5M 6F 6M E-AAC/M-CTT-58* BIN4.1 VIT151* E-AAC/M-CAA-181 E-AAC/M-CAT-167 E-AAC/M-CCT-128 P-CAG/M-CAA-93 BIN4.2 POR029. PIE002. VIT003 PIE228. PIE216 WAG016. PIE194 E-AAC/M-CCT-210 BIN4.3 PIE078 MSQ4 VIT043. PIE188 BIN4.4 PIE158. PIE176 PIE183. VIT018 E-AAG/M-CTA-150/5 0.0 BIN4.1 POR029 E-AAC/M-CTT BIN4.2 P-CCA/M-CAA PIE228. PIE216 BIN4.3 PIE088. VIT135 VIT151 MSQ E-AAC/M-CAA-543/ BIN4.4 B / BIN5.1 QrZAG39 E-AAC/M-CAC-199 E-AAG/M-CAC-332 BIN5.2 POR030. PIE246* PIE201. WAG017 FIR106*. WAG027 VIT036 BIN5.3 E-AAC/M-CAC-62 PIE100. VIT104 QpZAG58 R PIE246. PIE201 BIN5.1 WAG027. VIT023 FIR053. VIT036 QrZAG E-AAG/M-CTA E-AAG/M-CTT-65 FIR053. QRZAG73 BIN5.4 FIR035. VIT063. PIE013 GOT039. GOT037 E-AAC/M-CAA E-AAC/M-CTT-217 E-AAC/M-CAA WAG002. VIT141. VIT132 BIN6.1 VIT142. VIT094. WAG018 GOT040. FIR021. POR012*. PIE260. POR016. E-AAG/M-CTA POR021. POR024 VIT141. GOT063. PIE111. WAG002 VIT094. GOT040. FIR021. POR012 QrZAG5b WAG018. PIE260. FIR042.POR017 BIN6.1 POR034. POR018. FIR027. WAG041 MSQ13 WAG057. WAG059. VIT132. PIE035. FIR110. PIE125 BIN6.2 FIR110. PIE125 E-AAC/M-CTT BIN6.3 E-AAC/M-CAA-339 QpZAG64 E-AAC/M-CAA-75 BIN6.4 POR017. POR027 VIT089*. VIT121 B POR027*. VIT121. VIT138* MSQ BIN6.2 QpZAG E-AAC/M-CTT BIN5.5 POR014 E-AAC/M-CAA-149 QrZAG FIR024. VIT104 BIN5.2 VIT063 E-AAC/M-CCT PIE013*. WAG043 BIN5.3 GOT039. GOT037 QpZAG Durand et al., submitted.

16 Quercus and genomics Comparative mapping EST-SSRs in Quercus spp. HSLO3 0.0 *P13/M68_ *PIE022e 9.3 *PIE *POR *E37/M74_278a 25.7 *PIE *PIE *PIE *PORO *FIR *PIE *GOT021 3P x A4 0.0 PIE POR Po PIE PIE *P13/M66_ GOT *PIE075 *P14/M68_287a *ssrqrzag102 FIR PIE EVOLTREE, Jera 1.3 Bodenes et al., unpublished FIR *E38/M67_317a FIR004 linkage group 3 (male)

17 QTL mapping bud burst Quercus robur Castanea sativa Co-location of QTL in different species 17 Casasoli, M. et al. (2006) Genetics 172:

18 QTL mapping in oaks Quercus robur QTL mapping 18

19 QTL mapping in oaks 19 Q. robur x Q. robur subsp. slavonica, 387 full-sibs

20 QTL mapping stomatal density stomata 20 Q. robur x Q. robur subsp. slavonica, 387 full-sibs

21 QTL mapping stomatal density Trade-off between carbon fixation and water loss a b 100 μm 21 Gailing, O., Langenfeld-Heyser, R., Polle, A. & R. Finkeldey QTL loci affecting stomatal density and growth in a Quercus robur progeny: implications for the adaptation to changing environments. Global Change Biology 14:

22 QTL mapping stomatal density Aim of the study: Characterization of the genetic basis of stomatal density and plant growth under nonwaterstress conditions 22

23 QTL mapping stomatal density 23 Gailing, O., Langenfeld-Heyser, R., Polle, A. & R. Finkeldey QTL loci affecting stomatal density and growth in a Quercus robur progeny: implications for the adaptation to changing environments. Global Change Biology 14:

24 QTL mapping stomatal density Major-QTL for WUE (Brendel et al. 2007) LG11 PIE004 PIE033 QrZag18 P14-M E42-M74-82 PIE022f PIE040 PIE035 E37-M E41-M P13-M PIE036 PIE037 PIE275 PIE257 P14-M68-89 SD2 SD3 SD4 SD LG11 + PIE004 P13-M PIE033 QrZag18 PIE040 PIE035 E42-M74-90 E37-M E38-M74-96 P44-M PIE275 PIE257 E37-M SD2 SD3 SD4 SD5 24

25 Introduction adaptive traits Genotype - environment interaction Fitness Fitness 25 environment Mitchell-Olds, T. Willis, J.M. and Goldstein, D.B (2007) Nature Genetics 8:

26 QTL mapping stomatal density Perspectives Different water regimes Mapping of QTLs for carbon isotope composition (water use efficiency) Integration of candidate genes for this character (e.g. ERECTA) Comparative QTL mapping 26 Masle et al. (2005) The ERECTA gene regulates plant transpiration efficiency in Arabidopsis. Nature 436,

27 Adaptive genetic variation in oaks General Introduction Quercus and genomic resources Genetic mapping of drought related traits Species differentiation and local adaptation Outlook 27

28 Adaptive variation and species differentiation Quercus robur / Q. petraea Species distinction only by multiple characters (Kremer et al. 2002) petraea robur Generally low differentiation at molecular markers Kremer et al Strong differentiation at single markers (Scotti-Saintagne et al. 2004) Different adaptive potential 28 Scotti-Saintagne et al. 2004

29 Adaptive variation and species differentiation Outlier loci on genetic linkage maps! F ST between species Association with adaptive traits? 29 Scotti-Saintagne et al. (2004). Genetics 168:

30 Adaptive variation and species differentiation Quercus sensu strictu (Nixon 1993) Quercus robur L. Quercus petraea (Matt.) Liebl. Quercus frainetto Ten. Quercus pubescens Willd. Quercus cerris Bejan Forest (45 51 N, E), Carpathian mountains in westcentral Romania (Curtu, 2006) 30 Curtu, A.-L., Finkeldey R. & O. Gailing Comparative sequencing of a microsatellite locus reveals size homoplasy within and between European oak species (Quercus spp.). Plant Molecular Biology Reporter 22:

31 Adaptive variation and species differentiation Ecological requirements Quercus robur nutrient-rich wet soils Quercus petraea more acid, better drained soils Quercus frainetto xero-mesophile Quercus pubescens xerophile 31

32 Adaptive variation and species differentiation Q. frainetto Q. pubescens 32 Meusel H. and Jäger E.J. PSE 162:

33 Adaptive variation and species differentiation 33 Curtu, A.L., Gailing, O., Leinemann, L. & R. Finkeldey Genetic variation and differentiation within a natural community of five oak species (Quercus spp.). Plant Biology 9:

34 Adaptive variation and species differentiation Selection against hybrids? Generation F 1 forms Introgressive Total Sample size Percentage (%) Adult trees Offspring only for Q. petraea and Q. pubescens Adult trees Offspring

35 Adaptive variation and species differentiation Number of hybrids: low in adults / high in offspring Species are not randomly distributed in the forest Differentiation is generally low with exceptions 35

36 Adaptive variation and species differentiation Q. ellipsoidalis: xeric sites Q. rubra: mesic sites 36 Jensen et al

37 Adaptive variation and species differentiation Q. ellipsoidalis 37 Hipp and Weber, 2008

38 Adaptive variation and species differentiation Huron Mountain Reserve Ford Center, Baraga Plains km

39 Adaptive variation and species differentiation Ford Center, Baraga Plains Stand A Stand C Stand B 39

40 Adaptive variation and species differentiation Q. ellipsoidalis: xeric site Q. rubra: mesic sites 40 Quercus ellipsoidalis, Huron Mountain Reserve

41 41 Adaptive variation and species differentiation

42 Leaf morphological differentiation 42 according to Jensen et al. (1993). American Journal of Botany 80:

43 Leaf morphological differentiation Region Q. ellipsoidalis Q. rubra Huron Mountain Reserve (HMR) Ford Center (FC) A Ford Center (FC) B Ford Center (FC) C

44 Genetic markers Population Locus N N a N e H o H e F G0T021* Ford Center Huron Mountain Reserve PIE040 * quru-ga- 0E09** quru-ga- 1F07** quru-ga- 0C11** G0T021* PIE040 * quru-ga- 0E09** quru-ga- 1F07** quru-ga- 0C11** **nuclear SSRs (Aldrich et al. 2003) * EST-SSRs (Durand et al. unpublished)

45 45 Genetic differentiation

46 46 Acknowledgements

Impact of genetic variation in stomatal conductance on water use efficiency in Quercus robur. Oliver Brendel. INRA Nancy France

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