bioe 109 summer 2009 lecture 5- part ii selection, mutation & migration

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BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

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Page 1: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

BIOE 109Summer 2009

Lecture 5- Part IIselection, mutation & migration

Page 2: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Directional selection always maximizes mean population fitness!

Page 3: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Page 4: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of directional selection?

Page 5: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of directional selection?

Genotype: AA Aa aaFitness: 1 1 0.50

Here, selection is favoring a dominant allele.

Page 6: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Directional selection always maximizes mean population fitness!

Page 7: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of directional selection?

Genotype: AA Aa aaFitness: 0.4 0.4 1

Here, selection is favoring a recessive allele.

Page 8: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of balancing selection?

Page 9: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of balancing selection?

Genotype: AA Aa aaFitness: 1-s 1 1-t

Page 10: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of balancing selection?

Genotype: AA Aa aaFitness: 1-s 1 1-t

^Results in stable equilibrium point at p = t/(s + t)

Page 11: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of balancing selection?

Example: suppose s = 0.40 and t = 0.60

Page 12: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of balancing selection?

Example: suppose s = 0.40 and t = 0.60

Genotype: AA Aa aaFitness: 0.6 1 0.4

Page 13: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of balancing selection?

Example: suppose s = 0.40 and t = 0.60

Genotype: AA Aa aaFitness: 0.6 1 0.4

^Stable equilibrium point at p = t/(s + t)

Page 14: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Natural selection and mean population fitness

Question: How does w change during the process of balancing selection?

Example: suppose s = 0.40 and t = 0.60

Genotype: AA Aa aaFitness: 0.6 1 0.4

^Stable equilibrium point at p = t/(s + t)

= 0.60

Page 15: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Balancing selection maximizes mean population fitness!

Page 16: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

… but is only 0.75!

Balancing selection maximizes mean population fitness!

Page 17: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Conclusion:

Natural selection always acts to maximize mean population fitness

Page 18: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• Let p = frequency of A1 allele

Page 19: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• Let p = frequency of A1 allele

• Let q = frequency of A2 allele

Page 20: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• Let p = frequency of A1 allele

• Let q = frequency of A2 allele

• Let p = q = 0.50

Page 21: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• Let p = frequency of A allele

• Let q = frequency of a allele

• Let p = q = 0.50 u

A a

• Let u = 1 x 10-5 (and ignore v)

Page 22: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• Let p = frequency of A allele

• Let q = frequency of a allele

• Let p = q = 0.50 u

A1 a v

• Let u = 1 x 10-5 (and ignore v)

Page 23: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• denoting the change in a in one generation as q,

Page 24: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• denoting the change in a in one generation as q,

q = u x p

Page 25: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• denoting the change in a in one generation as q,

q = u x p

= (1 x 10-5)(0.5)

Page 26: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• denoting the change in a in one generation as q,

q = u x p

= (1 x 10-5)(0.5)

= 0.000005

Page 27: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• denoting the change in a in one generation as q,

q = u x p

= (1 x 10-5)(0.5)

= 0.000005

• the a allele has increased in frequency to 0.500005.

Page 28: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• denoting the change in a in one generation as q,

q = u x p

= (1 x 10-5)(0.5)

= 0.000005

• the a allele has increased in frequency to 0.500005.• it would take another 140,000 generations to reach 0.875

Page 29: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Mutation

• denoting the change in a in one generation as q,

q = u x p

= (1 x 10-5)(0.5)

= 0.000005

• the a allele has increased in frequency to 0.500005.• it would take another 140,000 generations to reach 0.875

Conclusion: The rate of change due to mutation pressure is extremely small!

Page 30: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Some comments on mutation

1. Mutation is the “raw material” that fuels all evolutionary change. 

Page 31: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Some comments on mutation

1. Mutation is the “raw material” that fuels all evolutionary change. 2. Mutations occur randomly! 

Page 32: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Some comments on mutation

1. Mutation is the “raw material” that fuels all evolutionary change. 2. Mutations occur randomly! 3. Mutations occur too infrequently to cause significant allele frequency change.

Page 33: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Some comments on mutation

1. Mutation is the “raw material” that fuels all evolutionary change. 2. Mutations occur randomly! 3. Mutations occur too infrequently to cause significant allele frequency change. 4. Most mutations are deleterious and experience “purifying selection”. 

Page 34: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Some comments on mutation

1. Mutation is the “raw material” that fuels all evolutionary change. 2. Mutations occur randomly! 3. Mutations occur too infrequently to cause significant allele frequency change. 4. Most mutations are deleterious and experience “purifying selection”. 5. A small (but unknown) proportion of mutations are beneficial and lead to adaptation.

Page 35: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Migration (gene flow)

Page 36: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Migration (gene flow)

• gene flow is simply the movement of genes among populations.

Page 37: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Migration (gene flow)

• gene flow is simply the movement of genes among populations.

• it can occur by the movement of gametes, or by the movement (and successful breeding) of individuals.

Page 38: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Migration (gene flow)

• gene flow is simply the movement of genes among populations.

• it can occur by the movement of gametes, or by the movement (and successful breeding) of individuals.

• its magnitude is determined by m

Page 39: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Migration (gene flow)

• gene flow is simply the movement of genes among populations.

• it can occur by the movement of gametes, or by the movement (and successful breeding) of individuals.

• its magnitude is determined by m

m = the proportion of genes entering a population in individuals (genes) immigrating from a different population.

Page 40: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Sewall Wright’s Continent-Island model

Page 41: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

Page 42: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = frequency of A1 allele on island

Page 43: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = frequency of A1 allele on island

• let pC = frequency of A1 allele on continent

Page 44: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = frequency of A1 allele on island

• let pC = frequency of A1 allele on continent

• let m = proportion of A1 alleles moving to island each generation

Page 45: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = frequency of A1 allele on island

• let pC = frequency of A1 allele on continent

• let m = proportion of A1 alleles moving to island each generation

• let p’I = frequency of A1 allele on island in the next generation

Page 46: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = frequency of A1 allele on island

• let pC = frequency of A1 allele on continent

• let m = proportion of A1 alleles moving to island each generation

• let p’I = frequency of A1 allele on island in the next generation

p’I = (1-m)(pI) + m(pc)

Page 47: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = frequency of A1 allele on island

• let pC = frequency of A1 allele on continent

• let m = proportion of A1 alleles moving to island each generation

• let p’I = frequency of A1 allele on island in the next generation

p’I = (1-m)(pI) + m(pc)

“resident” “immigrant”

Page 48: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = change in frequency of A1 allele on island from one generation to the next

Page 49: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = change in frequency of A1 allele on island from one generation to the next

pI = p’I - pI

Page 50: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = change in frequency of A1 allele on island from one generation to the next

pI = p’I - pI

= (1-m)(pI) + m(pc) - pI

Page 51: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

A simple model of migration

• let pI = change in frequency of A1 allele on island from one generation to the next

pI = p’I - pI

= (1-m)(pI) + m(pc) - pI

= m(pc – pI)

Page 52: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

An example:

Page 53: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

An example:

let pc = 0.75

Page 54: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

An example:

let pI = 0.25let pc = 0.75

Page 55: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

An example:

let pI = 0.25let pc = 0.75

let m = 0.10

Page 56: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

An example:

let pI = 0.25let pc = 0.75

let m = 0.10

let’s ignore back-migration

Page 57: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

An example:

let pI = 0.25let pc = 0.75

let m = 0.10

pI = m(pc – pI)

Page 58: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

An example:

let pI = 0.25let pc = 0.75

let m = 0.10

pI = m(pc – pI) = 0.10(0.75-0.25)

Page 59: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

An example:

let pI = 0.25let pc = 0.75

let m = 0.10

pI = m(pc – pI) = 0.10(0.75-0.25) = 0.050

Page 60: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Now:

pI = 0.30pc = 0.75

let m = 0.10

Page 61: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Now:

pI = 0.30pc = 0.75

let m = 0.10

pI = m(pc – pI) = 0.10(0.75-0.30) = 0.045

Page 62: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Conclusions

Page 63: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Conclusions

1. Gene flow can cause rapid evolutionary change.

Page 64: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Conclusions

1. Gene flow can cause rapid evolutionary change.

2. The long-term outcome will be the elimination of genetic differences between populations!

Page 65: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Conclusions

1. Gene flow can cause rapid evolutionary change.

2. The long-term outcome will be the elimination of genetic differences between populations!

But… natural selection act to oppose gene flow

Page 66: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Example: Lake Erie water snakes (Nerodia sipedon)

Page 67: BIOE 109 Summer 2009 Lecture 5- Part II selection, mutation & migration

Example: Lake Erie water snakes (Nerodia sipedon)