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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

32-1 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

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32-1Copyright 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

32-2Copyright 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

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

Species

• 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

32-4Copyright 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

32-5Copyright 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

32-6Copyright 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)

32-7Copyright 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

32-8Copyright 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)

32-9Copyright 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

32-10Copyright 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

32-11Copyright 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

32-12Copyright 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

p2 + 2pq + q2 = 1

– calculate frequencies from q2 (homozygous recessive)

32-13Copyright 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

32-14Copyright 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

32-15Copyright 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.)

32-16Copyright 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.)

32-17Copyright 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

32-18Copyright 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.)

32-19Copyright 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.)

32-20Copyright 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.)

32-21Copyright 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

32-22Copyright 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.)

32-23Copyright 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

32-24Copyright 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.)

32-25Copyright 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.)

32-26Copyright 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

32-27Copyright 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

32-28Copyright 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

32-29Copyright 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

32-30Copyright 2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint

Fig. 32.15: Models of speciation

(cont.)

(a)

32-31Copyright 2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint

Fig. 32.15: Models of speciation (cont.)

(cont.)

(b)

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

Fig. 32.15: Models of speciation (cont.)

(c)

32-33Copyright 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

32-34Copyright 2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint

Fig. 32.19: Origin of Heteronotia binoei

Copyright © Craig Moritz, University of Queensland(cont.)

32-35Copyright 2005 McGraw-Hill Australia Pty Ltd PPTs t/a Biology: An Australian focus 3e by Knox, Ladiges, Evans and Saint

Fig. 32.19: Origin of Heteronotia binoei (cont.)

Copyright © Craig Moritz, University of Queensland

32-36Copyright 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