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The Historical Development of Evolutionary Theory LIFS 691 Advanced Evolution What is evolution? • change in the distribution of phenotypes • change in genotypic frequencies • change in allelic frequences • to unfold, open, or unfold (Webster’s Dictionary) Mechanisms of Evolution • Natural Selection (including sexual selection) • Genetic Drift • Mutation • Gen Flow Natural Selection • defined as the differential survival and reproduction of heritable phenotypic variants • without heritability of phenotypes, the process is merely “selection” Lamarckism was alive and well in the late 19th century. • Darwin’s theory could not overcome problems created by the theory of Blending Inheritance Jean-Baptiste Lamarck (1744-1829) • Only about 1/3 of biologists at the time subscribed to Darwin’s theory of Evolution by Natural Selection; many evolutionary biologists still agreed with Lamarck’s ideas. Mendel’s garden Gregor Johann Mendel (1822-1884) “Discovery” of Mendel’s Work • Provided a theory of particulate inheritance, which seemingly overcame problems imposed by the theory of blending inheritance. Mendel’s peas • Unfortunately, many traits are not controlled by a single polymorphic locus; thus, Mendel’s laws did not have immediate generality. Many still doubted natural selection was the primary mechanism of evolution. The Birth of Mutation Theory • Noted sudden origins of seemingly novel phenotypes after many generations of stability • Hypothesized that biochemical changes were occurring, which became known as mutations. (Actually, these changes were the product of hybridization between species of plants, but this was unknown at the time.) Hugo de Vries (1848-1935) primroses Beginning of NeoDarwinian Theory Sewell Wright Sir Ronald Fisher (or Bakken’s lost twin?) An Alternative Synthesis • Some approached evolution theory from an understanding of genetics and development. Richard Goldschmidt • Goldschmidt’s ideas were made unpopular to protect selectionist theories from the threat of mutationist theories. Today, he is most remembered for the concept of a “Hopeful monster,” which was a only a very small part of his contribution to evolutionary biology. “Competing” Views in Evolution • Neutral Theory vs. Selectionist Theories • Adaptationist vs. Anti-adaptationist • Optimization vs. Quantitative Genetic Theories Methods for Examining Genes • Candidate Loci • Electrophoresis (mid-1960’s) • Sequencing (mid-1970’s) • Genomics (mid-1990’s) “In ‘nothing in biology makes sense except in the light of evolution’, nothing in evolution makes sense except in the light of population genetics.” Richard Lewontin, Harvard University “The science of population genetics is the auto mechanics of evolutionary biology.” Rama S. Singh, McMaster University Singh (2003), Genome 46: 938-942 Hardy-Weinberg Equilibrium Assumptions: • • • • • • No selection Infinite population size Random mating No mutation Simple genetic system (one locus, two alleles) Non-overlapping generations Hardy (1908), Science Hardy-Weinberg Equilibrium p = frequency of allele A q = (1-p) = frequency of allele a At equilibrium the frequencies of genotypes is as follows: AA p2 Aa 2p(1-p) aa (1-p)2 Hardy (1908), Science