Microevolution Biodiversity • Number and kinds of species on the earth Traditional Classification Prior to 1859, Aristotelian essentialism dominated – Classification and taxonomy treated as single activity – Species were eternal, immutable, and discrete – Morphological characters (‘essential traits’) used to describe a species – Observed variation among individuals treated as aberrations around a type (idealized) form for the species – Combined with Linnaean hierarchy (1700’s), this approach generated species classifications Hierarchical structure of life • • • • • • • • • Molecule Cell Tissue Organ Organ system Organism Population Community Ecosystem Fossils • Fossils of extinct animals related to those in the same area. • Succession of fossil types Pakistan's Siwalik Hills Geographic relationships of similar species (‘Sarawak Law’) There are similar species in different geographic places with similar climate. Hydromedusa maximiliani Emydura macquarii Disharmony of island biotas • the relative proportions of different taxa are not the same on islands as they are on the mainland. • represents a "grab-bag" rather than a broad range of all possible taxa --> those groups capable of long-distance dispersal Shared embryonic features • Comparative embryology shows how different adult structures of many animals have the same embryonic precursors. Vestigial organs in many species • Organs or structures remaining or surviving in a degenerate, atrophied, or imperfect condition or form. Common ‘design blueprints’ for structures with very different functions Shift in biological thought Radical shift in thinking during 19th century away from Aristotelian essentialism • • • • • Hutton & Lyell: uniformitarianism (old = current processes) Lamarck: inheritance of acquired traits Species were not eternal (many fossils of extinct forms discovered) Species were not immutable (e.g., domestication by humans through breeding designs) Variation in nature was not aberrant, but common and important for survival of individuals Elements of microevolution • Evolution: change in populations over time • Individuals develop but do not evolve • Darwin’s theory of evolution by natural selection: (descent with modification) – Individuals within a population vary for some trait – Some of these variations are heritable – Often, more offspring are produced than can survive (limited resources) – Survival and reproduction is not random: individuals with the most favorable variations are "naturally selected". Darwinian Evolution by Natural Selection Theory not accepted until after 70 yrs • 1860’s: world’s leading physicist Lord Kelvin = earth’s not old enough for gradual Darwinian evolution to account for the observed diversity – Based on wrong assumptions about the sources of energy heating the earth – Not known at that time that the sun is fission powered – Decay of radioactive elements unknown until 1900’s – Gross underestimate of Earth’s age Darwinian Evolution by Natural Selection Theory not accepted until after 70 yrs • Because Darwin had no idea how variability was generated in populations – Mutation was unknown – Critics: the amount of genetic variability was strictly limited and natural selection would halt when variability ran out. – 1900s: Thomas Morgan experimental work with fruit flies – Mutation generated new variation in every generation and in every trait. Darwinian Evolution by Natural Selection Darwin had no idea how variation was passed on from parent to offspring – Lack of a satisfactory theory of inheritance – Mendel’s theory of the particulate inheritance of genetic “factors” (i.e., genes) from 1865-1866 was unknown until 1900. – Mendel’s laws of segregation and independent assortment – 1953: Watson and Crick discovered the nature of DNA as the hereditary genetic material – Darwin adopted the theory of blending inheritance – Heritable variation will be lost each generation = natural selection could not operate – How discrete units of heredity (genes) produce continuous variation? – How discrete genes could account for speciation? Fig. 2-10, Ridley 2004, Evolution Anti-Darwinian ideas • Neo-Lamarckism – Inheritance of acquired traits – Refuted by experiments of Weismann • Orthogenesis – Straight-line inheritance: Spp evolve towards a fate. No proposed mechanism • Mutationist – Discrete new phenotypes derive from mutations = speciation – De Vries, Morgan, Goldschmidt Evolutionary (Modern) Synthesis • 1930’s-1940’s • Neo-Darwinism • Reconciled Darwin’s theory with genetics – Fisher, Haldane, Wright: Mutation + Nat. Sel. = Adaptive evolution – Chetverikov, Dobzhansky: genetic var & change in natural pops. • Other contributors – Mayr, Rensch, Stebbins, Simpson – Mutation + Recombination + Nat. Sel. within spp (µevol) = speciation & macroevolution Evolutionary (Modern) Synthesis • Foundations of modern evolutionary biology – – – – – – – Phenotype ≠ genotype. Genotype + environment = phenotype Acquired characteristics are not inherited Hereditary variation based on non-blending genes Polygenic inheritance = Continuous phenotypes Alleles: isostable forms of a gene w/ small to large phenotypic effect Mutation + recombination = genetic variation of natural populations Evolutionary change is a population process Evolutionary (Modern) Synthesis – – – – – – – – – Genetic drift & Natural selection = genotypic changes in populations Nat. Sec. can produce slight or large changes (adaptations) Nat. pops are genetically variable and can evol fast if env. changes Genetic differentiation of populations in ≠ geographic regions (adaptive = from Nat. Sel.): often polygenic and quantitative traits = evolution is gradual Populations have phenotypically different genotypes Species are distinct non-interbreeding “gene pools” Speciation = origin of >1 species from a single common ancestor (often by genetic differentiation of geographically isolated pops) Higher taxa arise by accumulation of small differences (gradual) Fossil record has gaps (incomplete) but shows gradual evolution Modern Evolutionary Biology – > 1950’s: Molecular biology – Molecular evolution, mutation, genetic variation, species differences, development, phylogenetics – Neutral theory of mol. evol. (Kimura) – > 1960’s: Evolutionary ecology – Ecology, behavior, reproductive biol, life history. – EvoDevo – Evolutionary Genomics Neutralist-Selectionist Debate – 1960’s to present There are four mechanisms for evolutionary change with debatable relative importance (mainly 1 vs 3) 1. Natural selection 2. Mutation (more a source of variation and not a mechanism for evolution per se) 3. Random genetic drift 4. Migration (gene flow) Organic Evolution • • • • Evolution: changes in a population’s gene pool (genetic makeup) over time. Natural selection: differential survival and reproduction of individuals that differ from one another in [heritable] characteristics (phenotypes). Survival of the fittest, not the strongest (really, of the good enough). Fitness: survival + reproductive output (via female and male functions).