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What evolutionary forces alter
Population Genetics Evolution depends upon mutation to create new alleles. Evolution occurs as a result of allele frequency changes within/among populations. What evolutionary forces alter allele frequencies?
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How do allele frequencies change
in a population from generation to generation?
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Allele frequencies in the gene pool: A: 12 / 20 = 0.6 a: 8 / 20 = 0.4 Alleles Combine to Yield Genotypic Frequencies
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Our mice grow-up and generate gametes for next generations gene pool
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Allele frequency across generations:
A General Single Locus, 2 Allele Model Freq A1 = p Freq A2 = q Genotypic frequencies are given by probability theory
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One locus, 2 Allele Model Genotype A1A1 A1A2 A2A2 Given:
In a diploid organism, there are two alleles for each locus. Therefore there are three possible genotypes: Genotype A1A1 A1A2 A2A2 Given: Frequency of allele A1 = p Frequency of allele A2 = 1 - p = q Then: Genotype A1A1 A1A2 A2A2 Frequency p2 2pq q2 A population that maintains such frequencies is said to be at Hardy-Weinberg Equilibrium
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Hardy-Weinberg Principle
When none of the evolutionary forces (selection, mutation, drift, migration, non-random mating) are operative: Allele frequencies in a population will not change, generation after generation. If allele frequencies are given by p and q, the genotype frequencies will be given by p2, 2pq, and q2
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Hardy-Weinberg Principle Depends Upon the Following Assumptions
There is no selection There is no mutation There is no migration There are no chance events 5. Individuals choose their mates at random
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The Outcome of Natural Selection Depends Upon:
Relationship between phenotype and fitness. (2) Relationship between phenotype and genotype. These determine the relationship between fitness and genotype. Outcome determines if there is evolution
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% survival to reproduction: A = 0.05 B = 0.10
12.2 Growth of 2 genotypes in an asexually reproducing population w/ nonoverlapping generations % survival to reproduction: A = 0.05 B = 0.10 Fecundity (eggs produced): A = 60 B = 40 C:\Figures\Chapter12\high-res\Evolution-Fig jpg Fitness A = 0.05 x 60 = 3 Fitness B = 0.01 x 40 = 4
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R = Per Capita Growth Rate = Represents Absolute Fitness
The rate of genetic change in a populations depends upon relative fitness: Relative Fitness of A = Absolute fitness A Highest Absolute Fitness WA = /4 = 0.75 Often by convention, fitness is expressed relative to the genotype with highest absolute fitness. Thus, WB = 4/4 = 1.0
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The fitness of a genotype is the average lifetime
contribution of individuals of that genotype to the population after one or more generations, measured at the same stage in the life history.
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12.3 Components of natural selection that may affect the fitness of a sexually reproducing organism
C:\Figures\Chapter12\high-res\Evolution-Fig jpg
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12.1(2) Modes of selection on a polymorphism consisting of two alleles at one locus
C:\Figures\Chapter12\high-res\Evolution-Fig jpg
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12.1(1) Modes of selection on a heritable quantitative character
C:\Figures\Chapter12\high-res\Evolution-Fig jpg
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Individuals may differ in fitness because of their underlying genotype
Incorporating Selection Individuals may differ in fitness because of their underlying genotype Genotype A1A1 A1A2 A2A2 Frequency p2 2pq q2 Fitness w11 w12 w22 Average fitness of the whole population: w = p2w11 + 2pqw12 + q2w22
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Given variable fitness, frequencies after selection:
Genotype A1A1 A1A2 A2A2 Freq p2 w11 2pq w12 q2 w22 w w w New allele frequencies after mating: p2 w11 pq w12 pq w12 q2w22 + + w w New Frequency of A1 New Frequency of A2
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Fitness: Probability that one’s genes will be represented
in future generations. Hard to measure. Often, fitness is indirectly measured: (e.g. survival probability given a particular genotype) WAA WAa Waa s Fitness is often stated in relative terms Selection coefficient gives the selection differential
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Persistent Selection Changes Allele Frequencies
Strength of selection is given by the magnitude of the selection differential
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Selection Experiments Show Changes in Allele Frequencies
HW Cavener and Clegg (1981) Food spiked with ethanol
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Selection can drive genotype frequencies
away from Hardy Weinberg Expectations
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Predicted change in allele frequencies at CCR5
High frequency (Europe) High selection/transmisson (Africa) Predicted change in allele frequencies at CCR5 High frequency (Europe) Low selection/transmisson (Europe) Low frequency (Europe) High selection/transmisson (Africa)
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What is the frequency of A1 in the next generation?
p2w11 + pqw12 pt + 1 = p2w11 + 2pqw12 + q2w22 What is the change in frequency of A1 per generation? Dp = pt pt = p / w (pw11 + qw12 - w ) With this equation we can substitute values for relative fitness and analyze various cases of selection.
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Gene Action Fitness Relationship A1A1 A1A2 A2A2 1+s 1 + s 1 Dominance
Recessivity A1A1 A1A2 A2A2 1 + s t Overdominance A1A1 A1A2 A2A2 1 + s t Underdominance
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Dominance Genotype A1A1 A1A2 A2A2 Fitness 1 + s s S = 0.01 A1
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Recessive Genotype A1A1 A1A2 A2A2 Fitness 1 + s S = 0.01 A1
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Evolution in lab populations of flour beetles support theoretical predictions. Dawson (1970)
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Overdominance/Heterozygote Superiority
Genotype A1A1 A1A2 A2A2 Fitness 1 + s t S = t = Stable equilibrium is reached A1 Genetic diversity is maintained
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Viable allele did not fix in the population
Mukai and Burdick 1958
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Underdominance Genotype A1A1 A1A2 A2A2 Fitness 1 + s t S = t = 0.02 Unstable equilibrium A1 A1 maybe fixed or lost from the population
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Frequency-Dependent Selection
Allele frequencies in a population remain near an equilibrium because selection favors the rarer allele. As a result, both alleles are maintained in the population.
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Frequency-Dependent Selection
Perissodus
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Incorporating Mutation
Mutation alone is a weak evolutionary force
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However, mutation and selection acting in concert
are a powerful evolutionary force
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