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CHAPTER 24 POPULATION GENETICS Prepared by Brenda Leady, University of Toledo Copyright (c) The McGraw-Hill Com

CHAPTER 24

POPULATION

GENETICS

Prepared by

Brenda Leady, University of Toledo

Copyright (c) The McGraw-Hill Com

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Population genetics

Study of genes and genotypes in a population

Want to know extent of genetic variation, why it exists and how it changes over time

Helps us understand how genetic variation is related to phenotypic variation

Gene pool

All of the genes in a population Study genetic variation within the gene pool and how variation changes from one generation to the next Emphasis is often on variation in alleles between members of a population

Population

Group of individuals of the same species that can interbreed with one another

Some species occupy a wide geographic range and are divided into discrete populations

Genes in Natural Populations Are Usually Polymorphic  Polymorphism – many traits display variation within a

Genes in Natural Populations Are Usually Polymorphic

Polymorphism – many traits display variation within a population

Due to 2 or more alleles that influence phenotype

Polymorphic gene- 2 or more alleles Monomorphic – predominantly single allele Single nucleotide polymorphism (SNPs)

Smallest type of genetic change in a gene

Most common – 90% of variation in human gene sequences

Large, healthy populations exhibit a high level of genetic diversity

Raw material for evolution

Allele and genotype frequencies

Related but distinct calculations

Allele and genotype frequencies  Related but distinct calculations 7

Example

49 red-flowered RR 42 pink-flowered Rr 9 white-flowered rr

Allele frequency of r

Example  49 red-flowered RR  42 pink-flowered Rr  9 white-flowered rr  Allele frequency

1.0 - 0.3 = 0.7 frequency of R

Genotype frequency of rr

Example  49 red-flowered RR  42 pink-flowered Rr  9 white-flowered rr  Allele frequency

Hardy-Weinberg equation

Relates allele and genotype frequencies under certain conditions

Hardy-Weinberg equation  Relates allele and genotype frequencies under certain conditions 9

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Conditions…

The population is so large that allele frequencies do not change due to random sampling error

The members of the population mate with each other without regard to their phenotypes and genotypes

No migration occurs between different populations

No survival or reproductive advantage exists for any of the genotypes—in other words, no natural selection occurs

No new mutations occur

In reality, no population meets these conditions

If frequencies are not in equilibrium, an evolutionary mechanism is at work

Microevolution

Changes in a population’s gene pool from generation to generation

Change because…

Introduce new genetic variation (mutations, gene duplication, exon shuffling, horizontal gene transfer)

Population will not evolve with mutations as the only source

Evolutionary mechanisms that alter the prevalence of an allele or genotype (natural selection, random genetic drift, migration, nonrandom mating)

Potential for widespread genetic change

Selective survival of genotypes that confer greater reproductive success Natural selection acts on

Characteristics with a survival advantage

Make organisms better adapted, more likely to survive, greater chance to reproduce

Favors individuals that produce viable offspring

Modern description of natural selection

1. Allelic variation arises from random mutations that may alter the function of the protein.

  • 2. Some alleles may encode proteins that enhance an individual’s survival or reproductive success compared to that of other members of the population

  • 3. Individuals with beneficial alleles are more likely to survive and contribute their alleles to the gene pool of the next generation

  • 4. Over the course of many generations, allele frequencies of many different genes may change through natural selection, thereby significantly altering the characteristics of a population

Net result of natural selection is a population that is better adapted to its environment and/or more successful at reproduction.

Darwinian fitness

Relative likelihood that a genotype will contribute to the gene pool of the next generation as compared with other genotypes

Measure of reproductive success Hypothetical gene with alleles A and a

AA, Aa, aa

Suppose average reproductive successes are…

AA 5 offspring Aa 4 offspring Aa 1 offspring

Fitness is W and maximum is 1.0 for genotype with highest reproductive ability

Fitness of AA: W AA = 5/5 = 1.0 Fitness of Aa: W Aa = 4/5 = 0.8 Fitness of aa: W aa = 1/5 = 0.2

Mean fitness of population

Average reproductive success of members of a population

As individuals with higher fitness values become more prevalent, natural selection increases the mean fitness of the population

Natural selection patterns

Directional selection Stabilizing selection Disruptive selection Balancing selection

Directional selection

Favors individuals at one extreme of a phenotypic distribution that have greater reproductive success in a particular environment

Initiators

New favored allele introduced Prolonged environmental change

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Stabilizing selection

Favors the survival of individuals with intermediate phenotypes

Extreme values of a trait are selected against

Clutch size

Too many eggs and offspring die due to lack of care and food

Too few eggs does not contribute enough to next generation

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Disruptive selection

Favors the survival of two or more different genotypes that produce different phenotypes

Likely to occur in populations that occupy diverse environments

Members of the populations can freely interbreed

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Balancing selection

Maintains genetic diversity Balanced polymorphism

Two or more alleles are kept in balance, and therefore are maintained in a population over the course of many generations

2 common ways

For a single gene, heterozygote favored

Heterozygote advantage – H S allele

Negative frequency-dependent selection

Rare individuals have a higher fitness

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Sexual selection

Form of natural selection

Directed at certain traits of sexually reproducing species that make it more likely for individuals to find or choose a mate and/or engage in successful mating

In many species, affects male characteristics more intensely than it does female

Intrasexual selection

Between members of the same sex

Horns in male sheep, antlers in male moose, male fiddler crab enlarged claws

Males directly compete for mating opportunities or territories

Intersexual selection

Between members of the opposite sex Female choice Often results in showy characteristics for males Cryptic female choice

Genital tract or egg selects against genetically related sperm

Inhibits inbreeding

Explains traits that decrease survival but increase reproductive success

Male guppy (Poecilia reticulata) is brightly colored compared to the female

Females prefer brightly colored males

In places with few predators, the males tend to be brightly colored

In places where predators are abundant, brightly colored males are less plentiful because they are subject to predation

Relative abundance of brightly and dully colored males depends on the balance between sexual selection, which favors bright coloring, and escape from predation, which favors dull coloring

Seehausen and van Alphen Found That Male Coloration in African Cichlids Is Subject to Female Choice

Seehausen and van Alphen Found That Male Coloration in African Cichlids Is Subject to Female Choice

Cichlidae have over 3,000 species

More different species that any other vertebrate species

Complex mating and brood care

Female play important role in choosing males with particular characteristics

Pundamilia pundamilia and Pundamilia nyererei

In some locations, they do not readily interbreed and behave like two distinct biological species

In other places they behave like a single interbreeding species with two color morphs

They can interbreed to produce viable offspring

 Hypothesized that females choose males for mates based on male’s coloration  Male were in

Hypothesized that females choose males for mates based on male’s coloration

Male were in glass enclosures to avoid direct competition

Goal to determine which of 2 males a female would prefer

Females’ preference for males dramatically different under different lights

Mating preference lost under monochromatic light

Sexual selection followed a diversifying mechanism

Random genetic drift

Changes allelic frequency due to random sampling error Random events unrelated to fitness Favors either loss or fixation of an allele

Frequency reaches 0% or 100%

Faster in smaller populations

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Bottleneck

Population reduced dramatically and then rebuilds

Randomly eliminated members without regard to genotype

Surviving members may have allele frequencies different from original population

Allele frequencies can drift substantially when population is small

New population likely to have less genetic variation

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Founder effect

Small group of individuals separates from a larger population and establishes a new colony

Relatively small founding population expected to have less genetic variation than original population

Allele frequencies in founding population may differ markedly from original population

Neutral theory of evolution

Non-Darwinian evolution Neutral variation

Much of the variation seen in natural populations is caused by genetic drift

Does not preferentially select for any particular allele

Most genetic variation is due to the accumulation of neutral mutations that have attained high frequencies due to genetic drift

Neutral mutations do not affect the phenotype so they are not acted upon by natural selection

Main idea is that much of the modern variation in gene sequences is explained by neutral variation rather than adaptive variation

Sequencing data supports this idea

Nucleotide substitutions much more likely in 3 rd base of codon (usually don’t change amino acid) than 1 st or 2 nd (usually does change amino acid)

Changing the amino acid is usually harmful to the coded protein

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Selectionists oppose the neutralist theory

Neutralists argue that most genetic variation arises from neutral genetic mutations and genetic drift

Selectionists argue that beneficial mutations and natural selection are primarily responsible

Both accept that genetic drift and natural selection both play key roles in evolution

Migration

Gene flow occurs when individuals migrate between populations having different allele frequencies

Migration tends to reduce differences in allele frequencies between the 2 populations

Tends to enhance genetic diversity within a population

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Nonrandom mating

One of the conditions required to establish the Hardy-Weinberg equilibrium is random mating

Individuals choose their mates irrespective of their genotypes and phenotypes

Forms of nonrandom mating

Assortative/disassortative Inbreeding

Assortative mating

Individuals with similar phenotypes are more likely to mate

Increases the proportion of homozygotes

Disassortative mating

Dissimilar phenotypes mate preferentially Favors heterozygosity

Inbreeding

Choice of mate based on genetic history

Does not favor any particular allele but it does increase the likelihood the individual will be homozygous

May have negative consequences with regard to recessive alleles

Lower mean fitness of a population if homozygous offspring have a lower fitness value

Inbreeding depression

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