BIOE 109 Summer 2009 Lecture 6- Part I Microevolution – Random genetic drift

Random genetic drift Definition: random changes in the frequencies of neutral alleles from generation to generation caused by “accidents of sampling”

Random genetic drift Definition: random changes in the frequencies of neutral alleles from generation to generation caused by “accidents of sampling” Q: What is a neutral allele?

Random genetic drift Definition: random changes in the frequencies of neutral alleles from generation to generation caused by “accidents of sampling” Q: What is a neutral allele? A: A neutral allele has no effect on fitness.

Random genetic drift Definition: random changes in the frequencies of neutral alleles from generation to generation caused by “accidents of sampling” Q: What is a neutral allele? A: A neutral allele has no effect on fitness. Q: How can alleles be neutral?

How can alleles be neutral?

1. Mutations among very similar amino acids

How can alleles be neutral? 1. Mutations among very similar amino acids (leu)  (val) CUC GUC

How can alleles be neutral? 1. Mutations among very similar amino acids (leu)  (val) CUC GUC 2. Mutations involving silent (synonymous) changes

How can alleles be neutral? 1. Mutations among very similar amino acids (leu)  (val) CUC GUC 2. Mutations involving silent (synonymous) changes (leu)  (leu) CUC CUU

How can alleles be neutral? 1. Mutations among very similar amino acids (leu)  (val) CUC GUC 2. Mutations involving silent (synonymous) changes (leu)  (leu) CUC CUU 3. Mutations in non-coding (nonfunctional) DNA

Codon bias shows that silent changes may not be neutral!

Random genetic drift Definition: random changes in the frequencies of neutral alleles from generation to generation caused by “accidents of sampling” What are RANDOM changes in frequencies? What is sampling error?

Some properties of random genetic drift

1. Magnitude inversely proportional to effective population size (N e ).

Some properties of random genetic drift 1. Magnitude inversely proportional to effective population size (N e ). 2. Ultimately results in loss of variation from natural populations.

Some properties of random genetic drift 1. Magnitude inversely proportional to effective population size (N e ). 2. Ultimately results in loss of variation from natural populations. 3. The probability of fixation of a neutral allele is equal to its frequency in the population.

Some properties of random genetic drift 1. Magnitude inversely proportional to effective population size (N e ). 2. Ultimately results in loss of variation from natural populations. 3. The probability of fixation of a neutral allele is equal to its frequency in the population. 4. Will cause isolated populations to diverge genetically.

Some properties of random genetic drift 1. Magnitude inversely proportional to effective population size (N e ). 2. Ultimately results in loss of variation from natural populations. 3. The probability of fixation of a neutral allele is equal to its frequency in the population. 4. Will cause isolated populations to diverge genetically. 5. Is accentuated during population bottlenecks and founder events.

What is effective population size?

in any one generation, N e is roughly equivalent to the number of breeding individuals in the population.

What is effective population size? in any one generation, N e is roughly equivalent to the number of breeding individuals in the population. this is equivalent to a contemporary effective size.

What is effective population size? in any one generation, N e is roughly equivalent to the number of breeding individuals in the population. this is equivalent to a contemporary effective size. N e is also strongly influenced by long-term history.

What is effective population size? in any one generation, N e is roughly equivalent to the number of breeding individuals in the population. this is equivalent to a contemporary effective size. N e is also strongly influenced by long-term history. this is equivalent to a species’ evolutionary effective size.

Factors affecting effective population size

1. Fluctuations in population size

Factors affecting effective population size 1. Fluctuations in population size - here, N e equals the harmonic mean of the actual population numbers:

Factors affecting effective population size 1. Fluctuations in population size - here, N e equals the harmonic mean of the actual population numbers: 1/N e = 1/t(1/N 1 + 1/N 2 + 1/N 3 + … 1/N t )

Factors affecting effective population size 1. Fluctuations in population size - here, N e equals the harmonic mean of the actual population numbers: 1/N e = 1/t(1/N 1 + 1/N 2 + 1/N 3 + … 1/N t ) Example: Over 3 generations, N = 2000, 30, 2000

Factors affecting effective population size 1. Fluctuations in population size - here, N e equals the harmonic mean of the actual population numbers: 1/N e = 1/t(1/N 1 + 1/N 2 + 1/N 3 + … 1/N t ) Example: Over 3 generations, N = 2000, 30, 2000 Arithmetic mean =

Factors affecting effective population size 1. Fluctuations in population size - here, N e is equal to the harmonic mean of the actual population numbers: 1/N e = 1/t(1/N 1 + 1/N 2 + 1/N 3 + … 1/N t ) Example: Over 3 generations, N = 2000, 30, 2000 Arithmetic mean = Harmonic mean = 87.4

Factors affecting effective population size 2. Unequal numbers of males and females

Factors affecting effective population size 2. Unequal numbers of males and females let N m = No. of males, N f = No. of females:

Factors affecting effective population size 2. Unequal numbers of males and females let N m = No. of males, N f = No. of females: N e = 4N m N f N m + N f

Factors affecting effective population size 2. Unequal numbers of males and females let N m = No. of males, N f = No. of females: N e = 4N m N f N m + N f Example: Breeding populations of northern elephant seals:

Factors affecting effective population size 2. Unequal numbers of males and females let N m = No. of males, N f = No. of females: N e = 4N m N f N m + N f Example: Breeding populations of northern elephant seals: 15 alpha males each controlling a harem of 20 females:

Factors affecting effective population size 2. Unequal numbers of males and females let N m = No. of males, N f = No. of females: N e = 4N m N f N m + N f Example: Breeding populations of northern elephant seals: 15 alpha males each controlling a harem of 20 females: census size (N) = 315

Factors affecting effective population size 2. Unequal numbers of males and females let N m = No. of males, N f = No. of females: N e = 4N m N f N m + N f Example: Breeding populations of northern elephant seals: 15 alpha males each controlling a harem of 20 females: census size (N) = 315 effective size (N e ) = 57.1

Factors affecting effective population size 3. Large variance in reproductive success reduces N e because a small number of individuals have a disproportional effect on the reproductive success of the population.

Factors affecting effective population size 4. Genetic Bottlenecks genetic bottlenecks refer to severe reductions in effective population size. Time NeNe

Genetic Bottlenecks genetic bottlenecks refer to severe reductions in effective population size.

Genetic Bottlenecks genetic bottlenecks refer to severe reductions in effective population size. Examples: the northern elephant seal (Mirounga angustirostis) and the cheetah (Acinonyx jubatus).

Founder effects occur when a new population is founded from a small number of individuals.

Founder effects occur when a new population is founded from a small number of individuals. Consequences:

Founder effects occur when a new population is founded from a small number of individuals. Consequences: 1. New population has a fraction of genetic variation present in the ancestral population.

Founder effects occur when a new population is founded from a small number of individuals. Consequences: 1. New population has a fraction of genetic variation present in the ancestral population. 2. Initial allele frequencies differ because of chance.

Founder effects occur when a new population is founded from a small number of individuals. Consequences: 1. New population has a fraction of genetic variation present in the ancestral population. 2. Initial allele frequencies differ because of chance. Example: the silvereye, Zosterops lateralis

Founder effects in the silvereye, Zosterops lateralis

Timing of island hopping…

… and reductions in allelic diversity

The interplay between drift, migration, and selection 1. Gene flow vs. drift

The interplay between drift, migration, and selection 1. Gene flow vs. drift random drift and gene flow act in opposition to each other!

The interplay between drift, migration, and selection 1. Gene flow vs. drift random drift and gene flow act in opposition to each other! random drift  allow genetic divergence of two populations.

The interplay between drift, migration, and selection 1. Gene flow vs. drift random drift and gene flow act in opposition to each other! random drift  allow genetic divergence of two populations. gene flow  prevents divergence.

The interplay between drift, migration, and selection 1. Gene flow vs. drift random drift and gene flow act in opposition to each other! random drift  allow genetic divergence of two populations. gene flow  prevents divergence. if N e m > 1, gene flow overrides drift and prevents divergence

The interplay between drift, migration, and selection 1. Gene flow vs. drift random drift and gene flow act in opposition to each other! random drift  allow genetic divergence of two populations. gene flow  prevent divergence. if N e m > 1, gene flow overrides drift and prevents divergence if N e m < 1, random drift can lead to genetic divergence.

The interplay between drift, migration, and selection 2. Gene flow and selection

The interplay between drift, migration, and selection 2. Gene flow and selection gene flow and selection usually act in opposition.

The interplay between drift, migration, and selection 2. Gene flow and selection gene flow and selection usually act in opposition. selection  favor different alleles in different populations (“local adaptation”).

The interplay between drift, migration, and selection 2. Gene flow and selection gene flow and selection usually act in opposition. selection  favor different alleles in different populations (“local adaptation”). gene flow  prevents adaptive divergence.

The interplay between drift, migration, and selection 2. Gene flow and selection gene flow and selection usually act in opposition. selection  favor different alleles in different populations (“local adaptation”). gene flow  prevents adaptive divergence. if m > s, gene flow can overpower local adaptation

The interplay between drift, migration, and selection 2. Gene flow and selection gene flow and selection usually act in opposition. selection  favor different alleles in different populations (“local adaptation”). gene flow  prevent adaptive divergence. if m > s, gene flow can overpower local adaptation if s > m, then selection can allow local adaptation

The interplay between drift, migration, and selection 3. Drift and selection

The interplay between drift, migration, and selection 3. Drift and selection - random genetic drift and natural selection act in opposition.

The interplay between drift, migration, and selection 3. Drift and selection - random genetic drift and natural selection act in opposition. selection  deterministic change in allele frequency

The interplay between drift, migration, and selection 3. Drift and selection - random genetic drift and natural selection act in opposition. selection  deterministic change in allele frequency random drift  random changes in allele frequency

The interplay between drift, migration, and selection 3. Drift and selection - random genetic drift and natural selection act in opposition. selection  deterministic change in allele frequency random drift  random changes in allele frequency if N e s > 10, selection controls the fate of the allele

The interplay between drift, migration, and selection 3. Drift and selection - random genetic drift and natural selection act in opposition. selection  deterministic change in allele frequency random drift  random changes in allele frequency if N e s > 10, selection controls the fate of the allele if N e s < 1, drift will overpower the effect of selection