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Partitioning Phenotypic Variability:
A Second View of Heritability (h2) VP = VG + Ve VP – Phenotypic variation VG – Genetic variation Ve – Unexplained variation
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Partitioning Phenotypic Variability:
A Second View of Heritability (h2) h2 = VG / VP
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Partitioning Phenotypic Variability:
A More Complete View Vp = VG + VE + VGxE + Cov(G,E) + Ve VE – Variation due to environmental factors: plastic response VGxE – Variation due to an interaction between genotype and the environment that influences phenotype
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Genotype by Environment Interactions
Response by a single genotype
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Genotype by Environment Interactions
Vg in high light Vg in low light
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Genotype by Environment Interactions
VE
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Genotype by Environment Interactions
VGxE – Degree to which lines are not parallel
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Partitioning Phenotypic Variability:
A More Complete View Vp = VG + VE + VGxE + Cov(G,E) + Ve Cov(G,E) – non-random association between genetic makeup and local environmental conditions
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Sources of Genetic Variation
Factors increasing variation Mutation 10-8 to mutations per base pair per generation 10-5 to 10-7 mutations per gene per generation One individual out of 10 per generation
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Sources of Genetic Variation
Factors increasing variation Mutation Migration Population #1 Population #2 Pollen Seeds
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Sources of Genetic Variation
Factors decreasing variation Natural selection Genetic drift in small populations (<1000) Loss of a allele Small population Large population p = frequency (allele A) q = frequency (allele a) p + q = 1 Loss of A allele
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Evidence of Selection in Natural Plant Populations
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Selection Among Populations
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The Common Garden Experiments of Clauson, Keck and Heiesy (1948)
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Differences in phenotype across a gradient: Yarrow (Achiella spp) as an example
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What is the source of variation?
Different species – genetic variation? Same species – phenotypic plasticity?
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Common Garden Experiment
Step #1: Obtain Plants from Source Populations Stanford – 100’ Timberline – 10,000’ Mather – 4600’
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Clones (e.g., piece of root)
Common Garden Experiment Step #2: Produce Clones Source Plant Clones (e.g., piece of root) Location #1 Location #2
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Clones (e.g., piece of root)
Common Garden Experiment Step #3: Plant clones in common gardens Source Plant Clones (e.g., piece of root) Common Gardens Location #1 Location #1 Location #2 Location #2
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Stanford Common Garden
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Mather Common Garden
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Timberline Common Garden
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Interpretation of Results: Pure Plastic Response
Source Plant Clones Common Gardens ? Location #1 Location #1 ? Location #2 Location #2
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Interpretation of Results: Pure Genetic Response
Source Plant Clones Common Gardens ? Location #1 Location #1 ? Location #2 Location #2
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Experimental Outcome: Growth of Mather Achiella Clones
Genetic response Plastic response
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Potentilla glandulosa
A Second Example Potentilla glandulosa Copyright © by Jane Strong and Tom Chester
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Lowland Plant Lowland Ecotype
©Brother Alfred Brousseau, St. Mary's College
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Montane Plant Potentilla glandulosa ssp ashlandica
©Brother Alfred Brousseau, St. Mary's College
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Experimental Outcome: Growth of Potentilla Clones
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What is the relationship between these organisms?
Interpretation Part I Not a pure plastic response Not a pure genetic response What is the relationship between these organisms? Separate experiments show that crosses between different source populations produce viable offspring
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Interpretation Part II
These are not different species What then are they?
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Ecotypes the middle ground
Genetically distinct organisms Phenotypically distinct in terms of Morphology Physiology Phenology Occur in distinct habitats Differences can be traced to ecological differences in home habitat Plants are potentially interfertile (i.e., same biologicial species)
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An Interpretation Individuals or Ecotypes
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Selection Within a Population
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Purple loosestrife (Lythrum salicaria): an aggressive invasive species
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Purple Loosestrife and Tristyly
Three flower types (morphs) ♀ Pistal positions differ ♂ Stigma positions differ Pollination patterns No self pollination Each morph can pollinate the other two morphs Less frequent morphs have higher fitness
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Impact of Frequency-Dependent Selection on
Invading Populations of Purple Loosestrife Study system with 24 newly invaded sites censused over a 5 year period Low evenness during year zero Evenness predicted to increase due to frequency dependent selection among morphs No change line (y=x) Prediction is met, indicating a change in population due to natural selection
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Selection At a Global Scale
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Convergent Evolution Example #1: Desert plants
Euphorbiaceae: Africa Cactaceae: N. America Example #2: Alpine plants Campanulaceae: Africa Asteraceae: S. America
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Life Histories and Tradeoffs
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Key Stages in the Life-History of a Plant
Seed Maturation Growth Dispersal Flowering seed phase Dormancy Pollination Germination
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The Ideal Plant Grow large rapidly Live forever
Reproduce early and often
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Impact of Limiting Resources
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General Scheme of Resource Allocation
Reproduction Pollen Nectar Ovules Seeds Growth Leaves Stems Roots Rhizomes Maintenance Structural support Storage Defenses Basal metabolism 3 2 1 General order in which resources are used
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