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Zoogeografia Dr. Valerio Ketmaier [email protected] Variation The synthetic theory of evolution Nothing in Biology Makes Sense Except in the Light of Evolution Dobzhansky T (1973) The American Biology Teacher 35:125-129 Evolution evolvere [lat.]: develop based on something already existing Biological (or organic) evolution is change in the properties of populations of organisms or groups of such populations, over the course of generations. Futuyma, 1998 Evolution can be precisely defined as any change in the frequency of alleles within a gene pool from one generation to the next. Curtis & Barnes, 1989 Preconditions for evolution • Inherited similarity • Variation • Selection Evolution of Influenza A virus in New York State 413 complete genomes, ML Changing environment => • inherited similarity • (unpredictable) variation • selection New York: 1997–1998, orange; 1998–1999, yellow; 1999–2000, light green; 2001–2002, dark green; 2002–2003, light blue; 2003–2004, dark blue; 2004–2005, purple; global background, red. Nelson et al. 2006, PLOS Pathogens Principles of evolution Evolution is a historical process => inheritance No design in variation => mutations unpredictable Selection occurs by differential survival/reproduction of different genotypes Selection leads to adaptation Evolution as a historical process Constraints by previous adaptations Barton et al. 2007 No design in variation Escherichia coli under stressful freezing, thawing & growth (FTG) Insertions/deletions in cardiolipin synth. => cls down, phosphatidilglycerol up => increased membrane fluidity No predictable course of adaptation Insertions near universal stress proteins => uspB down, uspA unaffected => uspB possibly related to membrane fluidity Sleight et al. 2008, Genetics Selection occurs by differential survival/ reproduction of different genotypes Experimental evolution of learning ability in Drosophila Circles: even numbered generations Squares: odd numbered generations Triangles: control populations Higher learning ability causes higher reproduction („fitness“) Mery & Kawecki T. J. 2002, PNAS Selection leads to adaptation Local adaptation in Poecilia mexicana Morphometrics, 13 landmarks blue: non-sulfidic surface yellow: sulfidic surface Plasticity or orange (LA): non-sulfidic cave red (CA): sulfidic cave Inherited change Genetics, 10 microsatellites ? Tobler et al. 2008, Evolution Plasticity or inherited changes ? Phenotypic plasticity => morphological differences caused by the environment (heritability = 0) Differences purely genetic => morphological differences genetically determined (heritability = 1) Heritability: h2 = VGenotype/VPhenotype Phenotypic trait with H2=0 (rare) „Acquired“ lack of tail in mice August Weismann (1892): (from Kutschera 2006) Phenotypic trait with H2=1 (rare) Mendel experiments Traits based on a single gene or tightly linked genes, no environmental influence Common: • Traits based on multiple gene loci (QTLs) • Environment impacts traits What is synthesized in the synthetic theory of evolution? Evolutionary theory „meets“ Genetics Mendel 1822-1884 Darwin 1809-1882 De Vries 1848-1935 Haeckel 1834-1919 Morgan 1866-1945 Evolutionary theory Principles: • Phenotypic variation • Limited resources • Survival of the fittest (natural selection) Side notes: • No genetics • Focused on quantitative traits Mendelians Principles: • new species through single mutations • quantitative variation not heritable • only discrete variation relevant • natural selection is old-fashioned • mutations and selection are mutually exclusive Side note: • focused on single gene traits • quantitative genetics unknown Synthetic theory of evolution (origin 1930-1950) „Mutations create the variation, which selection acts upon“ Theodosius Dobzhansky (Russian geneticist; 1900-1975): • 1927 emigrated to USA • „Genetics and the origin of species“ (1937) • biological species concept Sir Julian Huxley (English biologist; 1887-1975): • „Evolution, the modern synthesis“ (1942) How did giraffes get a long neck? Okapi Creationism / Intelligent Design (ID): • Neck has been designed as an adaptation Giraffe How did giraffes get a long neck? Organic progression / Lamarckism Continuous stretching of neck Inheritance of acquired traits How did giraffes get a long neck? Synthetic theory of evolution „Mutations create the variation, which selection acts upon“ • Neck length is a phenotypically variable quantitative trait • Part of the variation in neck length is based on genetic variation • Individuals with longer necks have a higher fitness • This selection on phenotypes increases frequency Of QTL_alleles positively correlated with neck length Preconditions for evolution • Inherited similarity • Variation • Selection Genetic variation Mutations • point mutations • chromosome mutations • polyploidization Recombination • sexual recombination • other types of recombination Horizontal gene transfer • in prokaryotes • in eukaryotes Mutation rates in various organisms Rates in bacteriophages & prokaryotic microbes (Drake 1999, Annals of the New York Academy of Sciences) Mutation rates in various organisms Mutation rates in higher eukaryotes per base: highest for bacteriophages, lowest for higher eukaryotes per genome: similar in all microbes, higher in higher eukaryotes (Drake 1999, Annals of the New York Academy of Sciences) Evolution of mutation rates in prokaryotes Mutator alleles as fast track to adaptive evolution (Taddeei et al. 1997, Nature) Evolution of mutation rates in prokaryotes Mutation rates in Pseudomonas with/without phages Mutation rates with (black)/without (white) phages (line is ancestral mutation rate) (Pal et al. 2007, Nature) 67% phages extinct with mutator 33% phages extinct without mutator Mutations are caused by ionizing radiation Mutation rates in offsprings of Chernobyl accident liquidators Random Amplified Polymorphic DNA (RAPD) n New bands New bands (%) CB (internal control) 18 5 0.019 CA 18 28 0.104 External Control 28 1 0.002 1: mother 2: father (working as liquidator) 3: offspring conceived after exposure (CA) 4: offspring conceived before exposure (CB) 5: negative control 6: size standard (Weinberg et al. 2001, Proc. R. Soc. B) Mutations are caused by ionizing radiation Impacts of the Fukushima nuclear accident on the pale grass blue butterfly F1-May 2011 (Hiyami et al. 2012, Nature) Mutations are caused by ionizing radiation Impacts of the Fukushima nuclear accident on the pale grass blue butterfly Sept 2011 (Hiyami et al. 2012, Nature) Mutations are caused by ionizing radiation Impacts of the Fukushima nuclear accident on the pale grass blue butterfly F2 (Hiyami et al. 2012, Nature) Polyploidization Chromosome number in sturgeons • several polyploidizations • evolution towards functional diploidy Phylogeny based on cytochrome b black: polyploidizations (Ludwig et al. 2001, Genetics) Hybridisation & recombination after secondary contact Atlantic sea sturgeon (spawns at 13-18°C) European sturgeon (spawns at >20°C) Distribution of mitochondrial haplotypes (part of Fig. 1; Ludwig et al. 2002, Nature) Atlantic sturgeon: American females successful in Europe Baltic Sea Canada Gironde MtDNA (data from Ludwig et al. 2002) n=81 n=10 n=67 n=15 n=17 MHC n=27 American (red), European (blue) and other (white) genotypes in sturgeons (Tiedemann et al. 2006, Naturwissenschaften) Hybridisation & recombination after secondary contact Origin 2n "oxy" "stur" I1-I4 HE HO American sea sturgeon (A. oxyrinchus) St.Lawrence/ St. Johns (Canada) 54 52 1 1 0.07 0.04 European sturgeon (A. sturio) Gironde (France) 34 - 33 1 0.06 0.06 Baltic sturgeon (A. oxyrinchus x sturio) Baltic Sea (18th to 20th century) 30 6 10 14 0.79 0.73 (Tiedemann et al. 2006, Naturwissenschaften) Amazon mollies (Poecilia) P. latipinna-female Gulf of Mexico P. latipinna-male Gynogenesis Gynogenesis MM P. formosa-female (all-female species) P. mexicana Sexual reproduction M P P MM MP (Schlupp et al., 1994, Science) Lack of recombination in P. formosa Characteristics in P. formosa P. latipinna P. mexicana A Allele size; inferred origin HE HO n A Allele size HE HO n A Allele size HE HO (GT) 10 2 339,341* (l) 0.52 0.00 7 4 335-4 77 0.66 0.29 7 5 349-3 85 0.59 0.43 (GT)4CT(GT)6 1 199 (m) 0.00 0.00 7 9 207-2 63 0.92 0.86 7 1 199 0.00 0.00 (GA) 14 4 407*,411*, 413 (l) 393 (m) 0.67 1.00 7 5 403-4 17 0.76 0.57 7 1 393 0.00 0.00 (GA)16AA(GA)2 CA(GA)5 3 197 (l) 203,205* (m) 0.63 1.00 7 6 173-1 99 0.86 1.00 4 7 187-2 07 0.96 0.75 (AG) 12 2 146 (l) 154 (m) 0.52 1.00 7 1 146 0.00 0.00 7 3 148-1 54 0.58 0.29 (GA)13 2 118 (l) 126 (m) 0.52 1.00 7 3 118-12 2 0.54 0.71 7 5 126-1 36 0.79 0.71 (GA)22 3 122 (l) 144*,148 (m) 0.63 1.00 7 4 116-14 6 0.70 0.71 7 7 120-1 60 0.86 0.29 (TG)10 3 176,178 (l) 180 (m) 0.63 1.00 7 2 176, 178 0.49 0.43 7 4 174-1 82 0.63 0.29 Repeat sequence (Tiedemann et al. 2005, Molecular Ecology Notes) Bdelloidea (Rotifers) Only females known Unisexual reproduction: • no genetic exchange among lineages • genetic depletion • +/- identical offspring Alternation of generations in rotifers • amictic females => Parthenogenesis • mictic females • unfertilized eggs => males • fertilized eggs => dormant eggs => amictic females Horizontal gene transfer in viruses Barton et al. 2007 Horizontal gene transfer and the origin of eukaryotes Last Universal Common Ancestor (LUCA)