MOLECULAR ADAPTATION

1) Review of genetic variation in natural population

2) Pop. genetics and Molecular markers

3) Genetic variation at the protein level

4) Genetic variation at the DNA level

History
Evolutionary synthesis integrated Darwinian selection & Mendelian inheritance Emergence of population genetics field Modern synthesis: integrated discoveries from paleontology, taxonomy, biogeography & pop genetics

Tenets of the Modern Synthesis

1. Importance of populations as units of evolution 2. Natural selection as primary mechanism of evolutionary change 3. Gradualism explains large changes occuring over long periods of time

Components of populations
Population = localized group of individuals of a species which are more likely to interbred with each other than with other species Species= a group of populations whose members have the potential to interbred in nature

Population genetics & the gene pool

Interbreeding within populations reduces gene flow Gene flow = migration of fertile individuals between populations Gene pool= total aggregate of genes in a population at any given time

Population genetics and genetic equilibrium

A population whose allele & genotype frequencies do not change down generations is in genetic equilibrium Hardy-Weinberg theorem states that normal processes of sexual reproduction do not change allele & genotype frequencies

Hardy-Weinberg Theorem concepts

Recombination & segregation associated with sexual reproduction do not alter the gene pool of population Used to describe non-evolving populations; can identify presence of evolution (change in gene frequencies)
Mathematics of theorem can be used to determine expected allele & gene frequencies in populations

Hardy-Weinberg equilibrium is maintained if

1. Very large population size is maintained
2. Isolation from other populations is maintained 3. No net mutations alter the gene pool 4. Individuals in population mate randomly 5. Natural selection is not influencing population

Microevolution
Five causes of microevolution are: 1. Genetic drift; occurs in small populations 2. Gene flow; migration among populations 3. Mutation; alters allele frequencies

4. Nonrandom mating
5. Natural selection; differential survival & reproduction

Genetic drift
In small populations, chance events cause allele frequencies to change randomly by sampling errors Smaller the population the more important genetic drift Two types: I) bottleneck effect

2) Founder effect

Genetic drift (contd)

Bottleneck effect
Occurs when a population is reduced to very small size rapidly & then recovers
Small numbers of survivors not likely to be representative Reduction effects all individuals randomly Longer population remains small the greater is the effect

Genetic drift (contd) Founder events

Genetic drift following founding of a new population by a few individuals (islands, continents)
Smaller the founding population, less likely its gene pool will be representative of original population

Genotype of founders determines genetic makeup of population

If colony survives, random drift will affect allele frequencies until population becomes relatively large

Nonrandom mating
2 types: Inbreeding
Individuals mate with close neighbors rather than with more distant members of a population Effect is reduced frequency of heterozygous genotypes, increased frequency of homozygotes Inbreeding do not necessarily change allele frequencies but does change genotype frequencies

1) Inbreeding 2) Assortative mating

Assortative matings
Individuals mate with partners that are like themselves in certain phenotypic characters
Examples: 1. Toads pair up according to size relationships; males must be able to achieve amplexus with larger females 2. In humans tall women commonly pair with tall men 3. Humans tend to mate within races, thereby reinforcing differences among ethnic groups

Gene Flow
Tends to reduce between population differences, thereby homogenizing population genetic structure

Can counteract effects of natural selection, genetic drift in relatively small, isolated populations

Mutation
Can immediate change the gene pool by allele substitutions

Natural selection
Differential survival & reproduction among members of a population Produces adaptive evolution by accumulation of favorable genotypes in populations Adaptive evolution= produces new genetic variation and selection favoring relatively fit individuals

Mutation & Sexual recombination are processes that generate genetic variation
Mutations: rare, random but ultimate source of allelic variation Usually occur in somatic cells, so most are not heritable

Point mutations that alter protein enough to affect function are more often harmful than beneficial, not retained
Chromosomal mutations usually affect many gene loci & tend to disrupt developmental processes Mutation can produce adequate genetic variation in bacteria due to their very short generation times

Diploidy & Balanced Polymorphism preserve genetic variation

Diploidy: hides much genetic variation from selection by presence of recessive alleles in heterozygotes
Because recessive alleles not expressed in heterozygotes, less favored or harmful alleles may be retained in populations The more rare the recessive allele is, the greater its protection by heterozygosity Result of heterozygote protection is maintenance of large pool of alleles which may be beneficial if conditions change

Balanced polymorphism= the ability of natural selection to maintain diversity in a population Natural selection can preserve genetic variation due to heterozygote advantage Example: Sickle-cell anemia caused by recessive allele

Heterozygotes are resistant to malaria so have fitness advantage over both homozygotes in regions with malaria Distribution of recessive allele closely tied to malaria incidence

Effect of natural selection on characteristics in population varies depending on which phenotypes are most favored 1) Directional selection
Natural selection operates to shift underlying allele frequencies in a consistent direction. Directional selection occurs when their is a directional change in the environment or a new mutation produces a new phenotype that is more adaptive than existing phenotypes. There are many examples of directional selection -resistance of bacteria and other organisms to chemicals designed to control them (e.g. antibiotics, DDT)

2) Stabilizing selection
Selection operates against extreme phenotypes and favors intermediate ones.

Stabilizing selection often results when environmental conditions have been constant. Example: Human birth weight -- newborns with birth weights much more or much less than the average (ca. 7 lbs.) have increased mortality.

3) Disruptive selection
Selection against intermediate phenotypes in favor of extreme phenotypes Disruptive selection can result in the formation of genetically distinct "subpopulations/"

Sexual selection: form of natural selection which can result in pronounced differences between sexes
Sexual selection: selection for secondary sexual characteristics advantageous in acquiring mates Results in sexual dimorphism: male & female differ in size or other traits advantageous in mating competition or female choice Examples: body sizes, plumages, horns & antlers vary between sexes