32-1 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Chapter 32: Mechanisms of evolution

32-2 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Populations and their gene pools Population –group of individuals of the same species, usually occupying a defined habitat –over one or more generations, genes can be shared through entire range of population –asexual populations more difficult to define  characterised by similarities in phenotype Gene pool –sum of all genes in a population at a given time

32-3 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Species –many concepts proposed to define a species Biological species concept –groups of actually or potentially interbreeding natural populations which, under natural conditions, are reproductively isolated from other such groups (definition proposed by Mayr and others) Other species concepts emphasise different aspects

32-4 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Evolutionary change Microevolution –change in gene pools –natural selection  change due to impact of environment –genetic drift  random change Macroevolution –change at or above the level of species  speciation

32-5 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Genetic variation Genetic variation within populations drives evolution Variation arises from –mutation –recombination

32-6 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Mutation Spontaneous or induced change in DNA sequence –minor (e.g. nucleotide substitutions, deletions) –major (e.g. chromosome inversions, translocations) Effect of mutation is expressed in phenotype –neutral  no effect –disadvantageous  negative effect (reduces fitness) –advantageous  positive effect (increases fitness)

32-7 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Measuring genetic variation Methods of detecting and measuring genetic variations –phenotypic frequency –genotypic frequency –allele frequency

32-8 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Phenotypic frequency Some phenotypic traits allow a population to be characterised genetically –variation in phenotype is directly related to genotype –genetic markers Variations (polymorphisms) in phenotypic trait are controlled by different alleles –example: Rhesus (Rh) blood groups in humans  Rh+ (dominant)  Rh- (recessive)

32-9 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Genotypic frequency Where dominance exists, phenotypic frequency gives incomplete information about allele frequency –recessive allele gives rise to phenotype when individuals are homozygous –dominant allele gives rise to same phenotype whether individuals are homozygous or heterozygous Immunological tests identify allele combinations –distinguish between homozygous and heterozygous individuals

32-10 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Allele frequency Calculate frequencies with which certain alleles occur –proportion of total alleles –does not indicate combinations p + q = 1 where p and q are frequencies of each allele

32-11 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Hardy–Weinberg principle Model of relationship between allele and genotypic frequencies Phenotypic frequencies in a population tend to remain constant at equilibrium values that can be estimated from allele frequencies Hypothetical ideal population –equilibrium established after one generation

32-12 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Hardy–Weinberg equation Allows genotypic frequencies to be calculated from phenotypic frequencies –where dominance exists p 2 + 2pq + q 2 = 1 –calculate frequencies from q 2 (homozygous recessive)

32-13 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Assumptions of H–W Individuals mate at random The population is so large that it is not affected by genetic drift No mutation No migration No natural selection

32-14 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Microevolution H–W assumption: Individuals mate at random Random mating –trait has no effect on mate choice Assortative mating –trait has an effect on mate choice –phenotypically similar mates  positive assortative mating –phenotypically dissimilar mates  negative assortative mating

32-15 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Assumptions of H–W H–W assumption: The population is so large that it is not affected by genetic drift Chance of microevolutionary change in a population’s gene pool –some alleles are lost –other alleles become fixed In small populations, the chance of genetic drift is high (cont.)

32-16 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Assumptions of H–W (cont.) H–W assumption: No mutation Mutation introduces novel genetic variation and new alleles H–W assumption: No migration Migration can change composition of gene pools if different groups exhibit different allele frequencies (cont.)

32-17 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Assumptions of H–W (cont.) H–W assumption: No natural selection Natural selection acts on phenotypes Changes frequencies of genotypes that give rise to those phenotypes –fitter genotypes appear in greater proportion to less fit genotypes Moves allele frequencies away from equilibrium

32-18 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Natural selection 1.More individuals are produced each generation than can survive to have offspring themselves –some individuals die before they reach breeding age –what determines which die and which survive? 2.Variation exists between individuals in a population and some of this variation involves differences in fitness –fitness is an organism’s ability to survive (viability) and produce the next generation (fertility) –some individuals have greater fitness than others (cont.)

32-19 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Natural selection (cont.) 3.Fitter individuals make a relatively greater contribution to the next generation than the less fit individuals –fitter individuals produce more offspring than others 4.Differences in fitness between individuals are inherited –reproducing individuals pass on their characteristics to the next generation (cont.)

32-20 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Natural selection (cont.) Fitter individuals reproduce more successfully than less fit individuals Contribute proportionately more to the next generation Cumulative effect over generations –results in change in gene pool (cont.)

32-21 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Speciation and species’ concepts Speciation is the process by which new species are formed Defining the concept of species is complex and no single species’ concept is universally accepted –biological species’ concept –taxonomic or morphological species’ concept –recognition species’ concept –evolutionary species’ concept –cohesion species’ concept

32-22 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Species’ concepts Biological species’ concept –‘groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups’ –does not consider morphologically different species that can interbreed to produce hybrids or asexually- reproducing species Taxonomic species concept –species is defined by phenotypic distinctiveness –members of a species are morphologically alike –problems with convergence and mimicry (cont.)

32-23 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Species’ concepts (cont.) Recognition species’ concept –species are groups sharing a common mate recognition system –does not consider asexually reproducing species Evolutionary species’ concept –a species is a lineage of populations delineated by common ancestry and able to remain separate from other species Cohesion species’ concept –species have mechanisms for maintaining phenotypic similarity, including gene flow and developmental constraints

32-24 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Reproductive isolation All species concepts consider reproductive isolation (prevention of gene flow between species) to be an important factor in maintaining a species’ integrity Reproductive isolating mechanisms inhibit or prevent gene flow between species –ecological isolation –temporal isolation –ethological isolation –mechanical isolation –gametic isolation –postzygotic isolation (cont.)

32-25 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Reproductive isolation (cont.) Ecological isolation –species do not hybridise because they occupy different habitats Temporal isolation –species do not hybridise because they are not ready to mate at the same time –example: two plant species produce flowers at different times Ethological isolation –species do not recognise each other as potential mates because the courtship patterns differ between species –example: frogs of different species have different mating calls (cont.)

32-26 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Reproductive isolation (cont.) Mechanical isolation –species do not hybridise because reproductive structures differ –example: differences in pedipalps of male spiders Gametic isolation –species do not hybridise because sperm are inviable in female reproductive tract, do not recognise egg of other species or cannot enter egg Postzygotic isolation –species may produce hybrids but hybrids are inviable or are sterile

32-27 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Allopatric speciation Populations of ancestral species are split by geographical barrier –inhibits migration and disrupts gene flow between populations Divergence of populations due to natural selection and genetic drift Reproductive isolation may develop, so if populations were to be reunited, gene flow would not be re-established

32-28 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Sympatric speciation Sympatric speciation takes place without geographical separation of populations Disruption of gene flow occurs when groups of individuals become reproductively isolated from other members of the population Polyploidy is a mechanism by which this occurs –multiple sets of chromosomes –common in plants –also found in some animals

32-29 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Parapatric speciation Parapatric speciation occurs in adjacent populations Geographical ranges are in contact, but selection exerts different pressures on populations Eventually gene flow is interrupted and populations become reproductively isolated

32-30 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Fig : Models of speciation (cont.) (a)

32-31 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Fig : Models of speciation (cont.) (cont.) (b)

32-32 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Fig : Models of speciation (cont.) (c)

32-33 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Hybridisation Not all hybrids are inviable or sterile Hybrids between species may become parthenogenetic –produce young from eggs without fertilisation Avoids problems of chromosome pairing with mismatched sets of chromosomes –example: parthenogenetic triploid gecko Heteronotia binoei formed by two hybridisation events

32-34 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Fig : Origin of Heteronotia binoei Copyright © Craig Moritz, University of Queensland (cont.)

32-35 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Fig : Origin of Heteronotia binoei (cont.) Copyright © Craig Moritz, University of Queensland

32-36 Copyright  2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint Molecular evolution Molecular sequences have diverged from a common ancestral sequence Gene duplication and sequence divergence produces gene families Homologous genes are derived from a common ancestral gene –orthologous genes arise when a species with the ancestral gene splits into two species –paralogous genes arise by gene duplication in a line of descent