Genecology, Restoration, and Adapting to Climate Change Brad St.Clair USDA Forest Service, Pacific Northwest Research Station, Corvallis, OR Workshop on Restoring and Sustaining Western Landscapes: Interaction with Climate Change 2009 Ecological Society of America Annual Meeting, Albuquerque, NM Photo: Berta Youtie When considering ecosystem responses to climate change, it is important to consider genetics of adaptation and genetic variation in adaptive traits. Three reasons: 1. Plants are genetically adapted to their local climates – populations, not species, are the important biological unit of interest 2. Evolutionary adaptation will determine what happens to plant populations given climate change 3. Management of genetic variation may positively influence how plants respond and adapt to climate change Three questions: 1. How are plants adapted to their local climates? 2. Will plants naturally adapt to future climates? 3. What can we do to help plants adapt to future climates? Adaptation “Evolutionary” adaptation” The process whereby an organism becomes better suited to its environment 2. A characteristic of an organism whose form is the product of natural selection in a given environment 1. “Societal” Adaptation The adjustment of natural or human systems to new environments, which moderates harm or exploits opportunities (IPCC 2001) 1. How are plants adapted to their local climates? Evidence for adaptation: 1. 2. 3. Correlation between a character and environmental factors - the same form occurs in similar environments Comparisons of naturally-occurring variants in environments where they are hypothesized to function as adaptations Direct evidence from altering a character to see how it affects function in a given environment Evidence for adaptation comes from common garden (provenance) studies Genecology • The study of interspecific genetic variation of plants in relation to environments (Turresson 1923) • Seeks correlations between “plant type” and “habitat type” • Consistent correlations are taken to indicate adaptive significance Evidence for adaptation: Correlations between traits and source environments - Douglas-fir Genecology Study Grow families in a common environment Measure many adaptive traits Collect seed from many trees GIS Combination of Variables, Primarily Growth Douglas-Fir of Western OR and WA 3 2 1 0 Traits vs source environment -1 -2 -3 -4 -5 -10 -8 -6 -4 -2 0 2 December Minimum Temperature 4 6 Douglas-Fir Genecology Study Fall cold damage Bud-set r = 0.79 Qst = 0.68 r = 0.76 Qst = 0.29 Biomass Bud-burst r = 0.52 Qst = 0.13 r = 0.60 Qst = 0.21 1. Populations differ 2. Traits are correlated with source environments 3. Different traits show different patterns and scales of adaptation • Ultimately interested in survival, growth and reproduction Differences among species: distance needed to detect genetic differences in Northern Rockies (Rehfeldt 1994) Species Douglas-fir Elev. (m) Frost- Evolutionary free days mode Specialist 200 220 370 18 20 33 38 40 54 Intermediate Western redcedar 420 450 600 Western white pine none 90 Generalist Lodgepole pine Engelmann spruce Ponderosa pine Western larch Specialist Intermediate Intermediate Generalist Bluebunch Wheatgrass Genecology Study Collections from: 127 populations 2 families per population 5 cultivars Planted at 3 common garden test sites in 2006: Central Ferry, WA – warm, dry Lucky Peak Nursery, ID – cooler, dry Pullman, WA – cold, wet Measured for 22 traits: Size Phenology Morphology Correlations of individual traits with climate At Lucky Peak, 2008 data Dry Wt Inflor No. Heading Date Bloom Mature Date Date Plant Form Leaf Form Awns Jan Temp 0.12 0.11 0.02 0.17 0.15 -0.07 0.15 0.04 Aug Temp -0.09 -0.01 -0.19 0.07 0.13 0.07 0.28 -0.33 Spring Frost Date -0.03 -0.07 0.25 0.09 -0.07 0.04 -0.17 0.29 Fall Frost Date -0.03 0.04 -0.21 -0.02 0.06 0.00 0.16 -0.26 Annual Precip 0.22 0.01 0.10 -0.03 0.02 -0.23 -0.28 0.27 Aug Precip 0.22 0.05 0.08 -0.17 -0.13 -0.21 -0.23 0.27 In general, correlations with climate are not strong. Relative to trees, no strong local adaptation. • Larger plants are from wetter areas. • Plants with later heading dates are from areas with later spring frost and earlier fall frost • Plants with upright form are from areas with less precipitation. • Plants with narrow leaves are from hot, dry areas. • Plants with longer awns are from areas with later spring frost, earlier fall frosts, cooler summers, and more precipitation. California/Mountain Brome (Bromus carinatus) Two studies: 1. Blue Mountains - USFS R6/ARS Erickson/Johnson – 209 sources + Bromar at 2 sites 2. Willamette Valley – USFS Doede – 107 sources at 2 sites Conclusions: • Considerable population variation in growth, form, and phenology • Considerable differences between regions • But weakly associated with climate (r<0.5) Mather El. 1,400 m Native to three different elevations planted at three different elevations (Clausen, Keck & Hiesey 1940) Stanford El. 35 m Potentilla glandulosa from Timberline El. 3,030 m Evidence for adaptation: Comparisons of naturally-occurring variants in native environments – reciprocal transplant studies Stanford El. 35 m Mather Timberline El. 1,400 m El. 3,030 m Grown at Response functions derived from lodgepole pine provenance tests in British Columbia What about genetic variation at the level of DNA? Patterns of Adaptive Molecular Genetic Diversity Eckhart, Neale, et al. 2009 Neutral Genotype Phenotype Genotype - Non-neutral and associated with phenotype 2. Will plants naturally adapt to future climates? Three possibilities when environments change: 1. • Move Migrate to new habitats where suited 2. Stay • • Acclimate by modifying individuals to new environment (phenotypic plasticity) Evolve, primarily through natural selection of better suited individuals 3. Disappear • Extinction of local population What is the potential for migration? • Evidence for range expansion northward and up in elevation • Estimates of past migration rates vary – Davis and Shaw 2001: 200-400 m per yr – Aitken et al 2007: 100- 200 m per yr • But current rates of climate change would require 3-5 km per yr What is the potential for evolution through natural selection? Important factors include: • Phenotypic variation • Heritabilities • Intensity of selection/fecundity What is the potential for evolution through natural selection? Important factors include: • • • • Phenotypic variation Heritabilities Intensity of selection/fecundity Generation turnover Lenoir et al. 2008. A significant upward shift in plant species optimum elevation during the 20th century. Science 320: 1768-1771. Optimum elevation = maximum probability of presence Avg optimum elevation shift = 29 m per decade Much quicker for grassy species compared to woody species: grassy species: ~ 90 m shift between 1986-2005 compared to 1905-1985 woody species: ~30 m shift What is the potential for evolution through natural selection? Important factors include: • • • • • • • Phenotypic variation Heritabilities Intensity of selection/fecundity Generation turnover Levels of gene flow Mating system Structure of genetic variation/steepness of clines What is the potential for evolution through natural selection? Important factors include: • • • • • • • • • Phenotypic variation Heritabilities Intensity of selection/fecundity Generation turnover Levels of gene flow Mating system Structure of genetic variation Central vs peripheral populations Trailing edge vs leading edge What is the potential for evolution through natural selection? Important factors include: • • • • • • • • • • Phenotypic variation Heritabilities Intensity of selection/fecundity Generation turnover Levels of gene flow Mating system Structure of genetic variation Central vs peripheral populations Trailing edge vs leading edge Population size What is the potential for evolution through natural selection? Important factors include: • • • • • • • • • • • Phenotypic variation Heritabilities Intensity of selection/fecundity Generation turnover Levels of gene flow Mating system Structure of genetic variation Central vs peripheral populations Trailing edge vs leading edge Population size Biotic interactions What is the potential for evolution through natural selection? Important factors include: • • • • • • • • • • • • Phenotypic variation Heritabilities Intensity of selection/fecundity Generation turnover Levels of gene flow Mating system Structure of genetic variation Central vs peripheral populations Trailing edge vs leading edge Population size Biotic interactions Genetic correlations Species and populations most threatened by climate change: • • • • • Long-lived species Inbreeding species Small populations Fragmented, disjunct populations Populations at the trailing edge of climate change (southern, low elevation) • Species or populations with low genetic variation • Rare species • Populations threatened from habitat loss, fire, disease, insects What about phenotypic plasticity? • Phenotypic plasticity = the ability of an individual to change its characteristics (phenotype) in response to changes in the environment • Phenotypic plasticity is common in plants – Plants modify their phenology and growth in response to changes in environments • • • • Bud-set Bud-burst Flowering Acclimation to drought • However, patterns of genetic variation in adaptive characteristics associated with environmental variation suggest that phenotypic plasticity is insufficient – No single phenotypically plastic genotype is optimal in all environments 3. What can we do to help plants adapt to future climates? 3. What can we do to help plants adapt to future climates? 1. Deploy populations adapted to future climates Seed zones and breeding zones are used to ensure adaptability USDA Plant Hardiness Zones Randall and Berrang (2002) WA Dept Nat Resources Seed zones and breeding zones are largely delineated based on climate Randall (1996) OR Dept of Forestry Recently, efforts to develop seed zones for native restoration species Recommended four seed zones for the Blue Mountains, oriented eastwest But, seed zones and seed transfer rules developed for today‟s climates may not be appropriate for future climates. Assisted migration = Movement of species, provenances, or breeding populations to „new‟ sites where they are expected to be better adapted in the future Present 2030 2060 2090 Douglas-fir Seed zone #4 0-1000 ft Seed zone Figures by Lauren Magalska, OSU Present 2030 2060 2090 Douglas-fir Seed zone #4 0-1000 ft Seed zone Climate Figures by Lauren Magalska, OSU Present 2030 2060 Douglas-fir Seed zone #4 0-1000 ft Seed zone Climate 2090 Present 2030 2060 Douglas-fir Seed zone #4 0-1000 ft Seed zone Climate 2090 Present 2030 2060 Douglas-fir Seed zone #4 0-1000 ft Seed zone Climate 2090 Assisted Migration for Douglas-Fir Corresponds to a temperature change of 2.5 to 6.2ºC Climate change and assisted migration of lodgepole pine Lodgepole pine provenance test in B.C. Change in productivity (m 3/ha) 70 50 30 10 -10 — Optimized sources — Local sources -30 -50 -70 0 140 populations 60 test sites 2012 | 1 2038 | 2 2063 | 3 2088 | 4 2114 | 5 2139 6 MAT increase (°C) Wang et al. (2006) Global Change Biol. 12:2404. Local = productivity increased by 7% up to 1.5ºC (2030), but decreased above 2ºC. Optimal = productivity increased by 14-36%. Web-based Seed Transfer Decision-Support Tool • Will help select appropriate seedlots for planting or target proper markets with specific seedlots • Will work for multiple species using multiple climatic variables and various climate change scenarios Developed by Ron Beloin, Glenn Howe, Brad St.Clair Center for Forest Provenance Data Objectives: 1. Archive data from long-term provenance tests and seedling genecology tests 2. Make datasets available to researchers through the web May eventually include species other than trees 3. What can we do to help plants adapt to future climates? 1. Deploy populations adapted to future climates (assisted migration) 2. Promote natural migration and gene flow Avoid fragmentation and maintain corridors for gene flow But, • Seed migration may not be sufficient • Pollen flow may be limited by temperature-associated flowering phenology 3. What can we do to help plants adapt to future climates? 1. Deploy populations adapted to future climates (assisted migration) 2. Promote natural migration and gene flow 3. Enhance genetic diversity • • • Deploy provenance mixtures within sites or across landscapes Maintain diversity within provenances Establish genetic outposts for facilitating gene flow into adjacent native stands – small number may be effective 3. What can we do to help plants adapt to future climates? 1. Deploy populations adapted to future climates (assisted migration) 2. Promote natural migration and gene flow 3. Enhance genetic diversity 4. Conserve genetic diversity Conserving Genetic Diversity In situ conservation • Locate reserves in areas of high environmental and genetic diversity • Reduce disturbance probability and intensity – thinning, prescribed fire, fuels reduction, insect traps • Supplement existing variation with genetic outposts Ex situ conservation • Seed collections becomes more important with increasing threats to in situ reserves • Assisted migration (plantings) may also be considered a form of ex situ conservation Priorities for Conservation • • • • • • • Long-lived species Picea mexicana Rare species Rare, valuable variants Low genetic variation Small population sizes Fragmented, disjunct populations Populations at the trailing edge of climate change (southern, low elevation) • Threatened from habitat loss, fire, disease, insects 3. What can we do to help plants adapt to future climates? 1. 2. 3. 4. 5. Deploy populations adapted to future climates (assisted migration) Promote natural migration and gene flow Enhance genetic diversity Conserve genetic diversity Practice selection and breeding for adaptive characteristics Selection and Breeding Breed for drought hardiness and growth phenology Tests have been developed to assess cold and drought hardiness. Breeding per se is generally not needed – assisted migration already available. Breed for resistance or tolerance to pests A long-term, expensive, difficult prospect. Key pests are being addressed – Which others will become problematic? Biotech approaches may be the most effective (e.g., Bt insect toxins). Breed for broad adaptation Testing for drought hardiness Xylem cavitation 3-cm stem section Imposed drought Cavitated cell Summary 1. How are plants adapted to their local climates? 2. Will plants naturally adapt to future climates? 3. What can we do to help plants adapt to future climates? Acknowledgements • • • • • Glenn Howe – Oregon State University Daniel Chmura – Oregon State University RC Johnson – ARS, Pullman, WA Vicky Erickson – USFS, Region 6 Nancy Shaw – USFS Rocky Mtn Research Station Final model fit data well from 3 trials Critical CU minimal CU needed (below that level, forcing will not result in budburst) Optimal CU - level at which additional chilling will not reduce the minimal time need for forcing Date of Spring Budburst Observed (squares) or Predicted from Historical Records (points) Historical data Blue=Oly Pink=Corv Experimental data indicates that some winter warming will hasten date of spring budburst but more warming will delay it as chilling is not satisfied. • Population variation in phenology, crown size, fecundity associated with aridity and temperature (r=0.55-0.71) Lodgepole pine transfer functions for six sites in British Columbia