Natural Selection results in increase in one (or more) genotypes relative to other genotypes.

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1 Natural Selection results in increase in one (or more) genotypes relative to other genotypes. Fitness - The fitness of a genotype is the average per capita lifetime contribution of individuals of that genotype to the population after one or more generations, measured at the same stage in the life history. Example: parthenogenic weevils Portion of eggs of genotype A that survive to reproductive age is Each adult lays an average of 60 eggs. Fitness is (0.05) (60) = 3. This is the per capita replacement rate (R), also called the Absolute Fitness. For Genotype A, R = 3 For Genotype B, R = 4 Calculate Relative Fitness (w) WA = 3/4 = 0.75 WB = 4/4 = 1.00 Absolute fitness of the genotype divided by the absolute fitness of the most fit genotype.

2 Can also calculate average fitness of the population, w. For the parthenogenic weevils, frequency of A = p frequency of B = q w = (p) (w A ) + (q) (w B ) Rates of change due to selection depend on w, not R. R can describe the rate of increase in population size, not fitness. Another result of selection is that the average fitness, w, increases. Coefficient of Selection (s) Measures the intensity of selection against a genotype. s A = 1 - w i s for the most fit genotype = 0 Modes of Natural Selection Directional Selection for one extreme Stabilizing Selection for the average Disruptive Selection for both extremes

3 Directional Selection One Gene models Selection for the dominant phenotype Selection for one homozygote with incomplete dominance Selection for the recessive phenotype Selection for one homozygote with incomplete dominance Genotype AA AA A A Frequency p 2 2pq q 2 Fitness 1 1-(s/2) 1-s p = (1/2)spq (1-sq) Selection will continue as long as the allele A is present. Selection for the dominant phenotype Genotype AA AA A A Frequency p 2 2pq q 2 Fitness s p = spq 2 1-sq 2 p increases relative to q 2 rather than q.

4 Natural Selection predicts a low level of genetic variability in populations. However, populations tend to have relatively high levels of genetic variablity. Polymorphisms and Variation Why might we observe polymorphisms? Transitional polymorphisms - one allele is replacing another Balanced polymorphisms - stable frequency with both alleles present. Balancing selection heterozygote advantage Selection in a variable environment A fluctuating environment may favor different genotypes at different times. Temporal fluctuations in the environment may slow down fixation due to selection, but it will generally not preserve both alleles. Spatial variations, with a mosaic of resources is more likely to maintain polymorphisms (multiple-niche polymorphism).

5 Protein Polymorphism Protein polymorphism in Drosophila natural populations is about 42%. Extend this to all genes, then there are about 3000 polymorphic loci. This amount can not be maintained by balanced polymorphisms. Segregational Load Most individuals would be less than optimally fit at many loci, greatly reducing the average fitness of the population. Unrealistic that polymorphisms are maintained by balance. Two possible explanations Unrealistic for population to be at maximum possible fitness. Different alleles are selectively neutral. (Neutral Theory) Solutions: Combination of factors - neutral - selection in a heterogeneous environment - hitchhiking - selection for heterozygote

6 Hard Selection vs. soft selection Hard Selection basically, the organism is either equipped to survive in a particular environment, or it is not. Soft Selection organisms of different genotypes can survive, but they will compete with each other. Hard Selection can promote speciation. Soft Selection can promote multiple-niche polymorphisms. Frequency Dependent Selection Fitness depends on the frequency of a particular phenotype (genotype) in the population. Inverse frequency-dependent selection The rarer a genotype is in the population, the greater its fitness. This process can easily maintain polymorphisms. Positive frequency-dependent selection The fitness of a genotype increases as its frequency in the population increases.

7 Natural Selection with multiple loci Each locus affects different characters. These loci should evolve independently of each other. However, they may be linked ort here may be a functional relationship. The Adaptive Landscape

8 Sexual Selection Success in mating (and producing offspring) is based on features that are not necessarily associated with other aspects of fitness. Different strategies for males and females Females produce a limited number of gametes and therefore there is a high cost to inappropriate matings. Males produce a large number of gametes and there is little cost to inappropriate matings. Intrasexual selection Intersexual Selection Mate choice, usually female choice. Benefits of sexual selection

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