33-1 copyright 2010 mcgraw-hill australia pty ltd powerpoint slides to accompany biology: an...
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33-1Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Chapter 33: Mechanisms of evolution
33-2Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-3Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-4Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-5Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Genetic variation• Genetic variation within populations drives
evolution• Variation arises from
– mutation– recombination
33-6Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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)
33-7Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Measuring genetic variation• Methods of detecting and measuring genetic
variations– phenotypic frequency– genotypic frequency– allele frequency
33-8Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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)
33-9Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-10Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-11Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-12Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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)
33-13Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Hardy–Weinberg assumptions• Individuals mate at random• The population is so large that it is not affected by
genetic drift• No mutation• No migration• No natural selection
33-14Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Microevolution
Hardy–Weinberg 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
33-15Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Genetic drift
Hardy–Weinberg 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
33-16Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Mutation and migration
Hardy–Weinberg assumption: No mutation
• Mutation introduces novel genetic variation and new alleles
Hardy–Weinberg assumption: No migration
• Migration can change composition of gene pools if different groups exhibit different allele frequencies
33-17Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Natural selection
Hardy–Weinberg 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
Question 1:
For a trait controlled by a single locus with two alleles which shows incomplete dominance, how many phenotypes are possible?
a) One
b) Two
c) Three
d) Four
e) Five
33-18Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
33-19Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Natural selection (cont.)
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
33-20Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-21Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-22Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-23Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-24Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-25Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-26Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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 (behavioural) 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
33-27Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
Question 2
Which of the following is not an example of a pre-mating reproductive isolating mechanism?
a) Temporal isolation
b) Mechanical isolation
c) Use of different sex attractant pheromones in butterflies
d) Ecological isolation
e) Incompatible sperm and ova
33-28Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
33-29Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-30Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Fig. 33.17: Models of speciation
(a)
33-31Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-32Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Fig. 33.17: Models of speciation (cont.)
(b)
33-33Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-34Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Fig. 33.17: Models of speciation (cont.)
(c)
33-35Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
33-36Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Copyright © Craig Moritz, University of Queensland
Fig. 33.21: Origin of Heteronotia binoei
33-37Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Fig. 33.24: Origin of Heteronotia binoei (cont.)
Copyright © Craig Moritz, University of Queensland
33-38Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
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
Summary• Evolution is the process of genetic change in a
population over time • Evolution results in individuals who are able to
survive and reproduce in their environment well enough to produce viable offspring
• Biological evolution requires heritable variation and a force to act on this variation
• Natural selection occurs when individuals with heritable differences show differences in survival and reproduction.
33-39Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University
Summary (cont.)• Heritable traits from fitter individuals become more
numerous in a population over time• Natural selection may act differently on males and
females in the same population• Co-operation between individuals in a population
can increase their inclusive fitness• Speciation usually involves physical, ecological or
temporal isolation of populations, which diverge over time
33-40Copyright 2010 McGraw-Hill Australia Pty Ltd PowerPoint slides to accompany Biology: An Australian focus 4e by Knox, Ladiges, Evans and SaintSlides prepared by Karen Burke da Silva, Flinders University