Genitcal Theory of Natural Selection By Julie Sobolewski and Angela Rose Terms: Allele - one of several forms of the same gene Allele Frequency - the proportion of gene copies in a population that are at a given allele Evolution - origin of genetic variation by mutation or recombination, followed by changes in the frequencies of alleles and genotypes Fitness - the success of an entity in reproducing; average contribution of an allele or genotype to he next generation or to succeeding generations Natural Selection - differential reproductive success Reproductive Success - average lifetime contribution of individuals of a particular genotype to a population after one or more generations (generally includes average number of offspring produced and average number that survive) How Natural Selection Differs from Evolution: -evolution is a two step process the origin of variation followed by changes in the frequencies of alleles. -natural selection can only have an evolutionary effect when phenotypes differ in genotype - a feature can only evolve through natural selection if it has a positive contribution to the reproduction or survival of individuals that bear it. Beginnings of the Gentical Theory of Natural Selection: In 1930 R. A. Fisher published his work, The Genetical Theory of Natural Selection. -Population fitness - The consequences of natural selection depend on four things - the relationship between phenotype and fitness - the relationship between phenotype and genotype - a relationship between fitness and genotype -whether or not evolutionary change occurs Modes of Selection (Figure 12.1): Directional - when disadvantageous alleles are replaced by more advantageous alleles. -Coral Snakes Stabilizing- if an intermediate phenotype is fittest -Heterozygote advantage, each generation produces all three genotypes are produced which results in a stable equilibrium. - Sickle cell Diversifying - if two or more phenotypes are fitter than the intermediates between them -Multiple-niche polymorphism -Black-bellied seedcracker Advantageous Allele Frequency: Δp = (½ spq) ÷ (1 – sq) -A character state with even a minuscule advantage will be fixed by natural selection -Advantageous allele may initially be fairly common but is likely to be rare if it is a newly arisen mutation or if it was disadvantageous before an environmental change Food for Thought: -Do you think coloration rather than inconspicuousness is advantageous? Why or why not? (Figure 12.4) Deleterious Allele Frequency: up = (spq2) ÷ w solve for q q = (u ÷ s) ½ -Deleterious alleles can be hidden by heterozygous states but they can persist because they are repeatedly reintroduced either by mutation or by gene flow from other population in which they are favored by different environment -Going to move towards a stable equilibrium Inverse frequency-dependent selection: - the rarer a phenotype is, the greater its fitness Sex allocation: -defined as the distribution of resources to male vs. female reproductive function -Male to female ratio = 1:1 -Evolutionarily Stable Strategy (ESS) = stable equilibrium Food for Thought: -How could global warming affect sex allocation? -Are there any species where sex allocation my not apply as strongly, if at all? Inverse frequency-dependent selection does not just apply to sex allocation: -cichlids in Lake Tanganyika (dextral vs. sinistral) -Figure 12.16 How Strong is Natural Selection? -Peppered Moths used as example for very strong selection. -Kettlewell’s results- when moths were released in polluted area typical had lower survival rate. When released in unpolluted area typical had higher survival rate. -Well’s and Hooper claimed experiment and results were false because of methods used. -Further research used similar methods obtained some controversial results. Food for Thought: -Was Kettlewell right? If not would this weaken the case for evolution by natural selection? References: 1) Askew, R. R., Cook, L. M., Bishop, J. A. (1971). Atmospheric Pollution and Melanic Moths in Manchester and Its Environs. The Journal of Applied Ecology, 8 (1), 247-256. 2) Brakefield, Paul M., Liebert, Tony G. (2000). Evolutionary Dynamics of Declining Melanism in the Peppered Moth in the Netherlands. Proceedings: Biological Sciences, 267 (1456), 1953-1957. 3) Charnov, E. L. 1982. The Theory of Sex Allocation. Princeton University Press, NJ. 4) Cook, L. M., Dennis, R. L. H., Mani, G. S. (1999). Melanic Morph Frequency in the Peppered Moth in the Manchester Area. Proceedings: Biological Sciences, 266 (1416), 293-297. 6) Holdrege, Craig (1999). The Case of the Peppered Moth Illusion. Whole Earth, 1-6. 7) Lees, D. R., Creed, E. R. (1975). Industrial Melanism in Biston betularia: The Role of Selective Predation. The Journal of Animal Ecology, 44 (1), 67-83. 8) Wells, Jonathan (1999). Second Thoughts about Peppered Moths: This classical story of evolution by natural selection needs revising. The Scientist, 13, 13-16.