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Genetic erosion and pollution genetic and conservation consequences for European forest tree species François Lefèvre INRA, Avignon (France) Conclusion Non-equilibrium paradigm for in situ management of long lived species current population size genetic diversity adaptedness are not stable nor ideal quantities, just instantaneous values resulting from dynamic processes => think in terms of parameter changes and evolutions Conservation of genetic resources Monitoring evolutionary changes 1. General concepts and processes 2. Specific features for trees 3. Consequences for the management of genetic resources 1. General concepts and processes: genetic erosion Ne, effective population size a standardized measure of genetic evolution (decrease of diversity or increase of inbreeding) unit : size of a model population that follows the same genetic evolution Ne = 30 Ne = 7 Ne = => constant Ne => Ne decreases => no evolution stable evolution erosion accelerates 1. General concepts and processes: genetic erosion Departure from the model population that decreases Ne: Biology - true or partial dioecy - departure from panmixia (both directions) Demography - variation in population size across generations - variance (V) in mating success - generation overlap Environment - selection (Nunney, 2000) human impact 1. General concepts and processes: hybridization-introgression Hybridization (intra + interspecific) Natural Anthropogenic Type 1 Natural hybrid Type 2 Natural introgression Type 3 Natural hybrid zone F1 only (backcrosses) Hybrid swarm (sterile F1) Type 4 Hybridization no introgression (Allendorf et al, 2001) Type 5 Widespread introgression Type 6 Complete admixture 1. General concepts and processes: balanced effects demographic rescue increase diversity reduce inbreeding restoration risk of extinction if : (Lenormand, 2002) demographic swamping migration load reduce Ne extinction local taxon is not rare reproductive barriers are strong long generation time selfing or vegetative propagation differencial selection is enhanced 2. Specific features for trees 2. Specific features for trees : high diversity in spite of recent colonization history, high diversity is generally maintained within tree populations trees annual plants nb species mean nb pop. mean nb loci 196 9.2 18.1 226 18.1 16.2 He (total div. ) Hs (within-pop div. ) Gst (differentiation) 0.177 0.148 0.084 0.154 0.101 0.355 (Hamrick et al, 1992) 2. Specific features for trees : local adaptation... in spite of recent colonization history, local adaptation rapidly developped through selection … adaptive cline in sessile oak (Ducousso et al, 1996) 2. Specific features for trees : local adaptation & diversity in spite of recent colonization history, local adaptation rapidly developped through selection … and high diversity for adaptive traits is maintained within tree-pops h² AGCV (sA/m %) (Cornelius, 1993) height straightness morphol. traits wood density branch 0.28 0.28 0.23 0.50 0.26 11.10 16.25 14.73 5.34 16.30 (Ducousso, unpubl.) phenology 0.30 23.73 2. Specific features for trees : important gene flow the high diversity is explained by important seed and pollen dispersal, the long juvenile phase increases migration and reduces the founder effect expected at founding a new deme (Mariette, 2001) (Austerlitz et al, 2000) 2. Specific features for trees : temporal patterns erosion of diversity is delayed by temporal patterns of fecundity: annual stochasticity reduces the impact of selection, cumulative effect reduces drift Ne=31 Ne=83 700 Ne=57 Ne=85 7743 Ne=36 2210 (Krouchi et al, 2004) Ne=59 7822 Ne=92 Ne=76 398 584 7787 45719 2. Specific features for trees : adaptational lag adaptation is a dynamic process where selection is balanced by gene flow (migration load), interaction among species (Red Queen hypothesis) and temporal fluctuations of environment response function 142 pop Pinus contorta tested on 60 sites (Rehfeldt et al, 2001) 2. Specific features for trees : rapid adaptation tree populations have the potential for rapid adaptive changes ... % budset in Norway spruce date (Skroppa & Kohman, 1997) 2. Specific features for trees : climate change present tree populations and the next 1 or 2 generations will experience climate change within their own life : short term phenotypic plasticity adaptation long term migration 3. Consequences for management of genetic resources 3. Consequences for management: global perspective Habitat monitoring Environmental impact Population management environment landscape wild relative interactions domestic resource 3. Consequence for management : genetic erosion domestication (breeding and selection) : not a major threat for diversity in trees, except if low initial diversity population management : direct impact on demographic parameters (life cycle, dispersal, mating system, survival, mating success) should not affect the processes that maintain a high level of diversity within tree populations (Lefèvre, 2004) 3. Consequence for management : genetic pollution introduction, fragmentation and habitat disturbance influence hybridization risk assessment : assess hybridization, occurrence and frequency (difficult) ; estimate variation in frequency among cohorts ; estimate the relative fertility of local and hybrid types reduce the impact of hybridization if considered as a risk : eliminate hybrids and invading sp ; improve habitat to enhance competition ; (adapted from Wolf et al, 2001) 3. Consequence for management : genetic pollution hypoth.: large amount of transplanted material involved in the regeneration maladaptedness local pop. large Ne genetic diversity high low local pop. small Ne low low high high genetic diversity high low maladaptedness can be avoided but low diversity more likely higher impact when local population has small Ne (Lefèvre, 2004) Conclusion Non-equilibrium paradigm for in situ management of long lived species current population size genetic diversity adaptedness are not stable nor ideal quantities, just instantaneous values resulting from dynamic processes => think in terms of parameter changes and evolutions Conservation of genetic resources Monitoring evolutionary changes References Allendorf et al, 2001 TREE 16, 613-622 Austerlitz et al, 2000 Genetics 154, 1309-1321 Cornelius, 1993 Can J For Res 24, 372-379 Ducousso et al, 1996 Ann Sci For 53, 775-782 Hamrick et al, 1992 New Forests 6, 95-124 Krouchi et al, 2004 For Ecol Manage 197, 181-189 Lefèvre, 2004 For Ecol Manage 197, 257-271 Lenormand, 2002 TREE 17, 183-189 Nunney, 2000 Evol Biol 32, 179-194 Rehfeldt et al, 2001 Climatic Change 50, 355-376 Skroppa & Kohman, 1997 Forest Genetics 4, 171-177 Wolf et al, 2001 15, 1039-1053 Conserv Biol