One area lacking in Darwin’s analysis
1) Competition creating selective force
2) Variety of heritable traits
3) Selection of organisms to out-reproduce
others
4) Generations of over-reproduction to foster
population change
How is this variety
Modern evolutionary theory
Darwinian evolution + genetic analysis = modern
synthesis
Methodology: analyze a population’s gene pool, see if
its allele frequencies are changing over time
Population – all organisms of same species in an area
Gene pool – all of the genes of population considered
together
Allele frequency – percentage of a specific allele
Example of gene pool
Studying a population of rock pocket mice
Sandy-fur mice = RR (25 organisms) Sandy-fur mice = Rr (17 organisms) Black-fur mice = rr (3 organisms)
Total alleles = 25 x 2 + 17 x 2 + 3 x 2 = 90
R allele frequency = 25 x 2 + 17 x 1 = 67 / 90 =
74.4%
Allele frequencies and evolution
Update definition of evolution as change in
allele frequencies over generations
Ex: Lava flow creates black rock environment
Genotypes # organisms initially
# organisms
25 years later
RR
25
2
Rr
17
4
Calculate new allele frequency
Initially: R = 74.4% r = 25.6%
Genotypes # organisms initially
# organisms
25 years later
RR
25
2
Rr
17
4
rr
3
41
25 years later: Total alleles = (2 + 4 + 41) x 2
= 94
R = 2 x 2 + 4 x 1 = 8 / 94 = 8.5%
So what causes evolution?
More than just natural selection
First, let’s discuss when no change occurs =
Hardy – Weinberg equilibrium
Hardy and Weinberg create mathematical
Hardy – Weinberg equilibrium
Population allele frequencies will remainconstant if:
1.No gene flow (emigration or immigration)
2.No mutations occur
3.No natural selection is occurring
4.Population is large in number (infinitely large)
Hardy – Weinberg equilibrium
If 5 assumptions met, no evolution occurs
This also means that sexual reproduction itself
does NOT change population allele frequencies
Sexual reproduction does create genetic variety
in individuals BUT NOT in population
So what causes evolution?
Let’s break the 5 assumptions one at a time
1.No gene flow (emigration or immigration)
2.No mutations occur
3.No natural selection is occurring
4.Population is large in number (infinitely large)
Evolutionary force #1: gene flow
Entry of new individuals or exit of current
members
Initially: R = 74.4% r = 25.6%
Genotypes # mice in desert
# mice + 10 new black
fur mice
RR
25
25
Rr
17
17
rr
3
13
After gene flow: R = 60.9% r = 39.1%
So what causes evolution?
Let’s break the 5 assumptions one at a time
1.No gene flow (emigration or immigration)
2.No mutations occur
3.No natural selection is occurring
4.Population is large in number (infinitely large)
Evolutionary force #2: mutation
Causes very little change by itself
Example: new mutation at another gene locus
causes albino coloration in just one mouse
(updating Darwin)
Genotypes # organisms initially
# organisms
after mutation
event
AA
0
0
Aa
0
1
aa
45
47
Before:
A = 0%
a =
100%
After:
A = 1%
So what causes evolution?
Let’s break the 5 assumptions one at a time
1.No gene flow (emigration or immigration)
2.No mutations occur
3.No natural selection is occurring
4.Population is large in number (infinitely large)
Evolutionary force #3 – natural selection
The example we discussed before (mice
invading black volcanic rock territory)
Genotypes # organisms initially
# organisms
25 years later
RR
25
2
Rr
17
4
rr
3
41
(Darwin’s idea)
Before:
R = 74.4%
r = 25.6%
After:
Real life examples
1) Stabilizing selection – child birth weight
2) Directional selection – anteater snout
length
3) Disruptive selection – limpet shell
So what causes evolution?
Let’s break the 5 assumptions one at a time
1.No gene flow (emigration or immigration)
2.No mutations occur
3.No natural selection is occurring
4.Population is large in number (infinitely large)
Evolutionary force #4: Genetic drift
Before, we said sexual reproduction does
NOT change overall gene pool
This assumes a large population where actual
results more likely match predictions
R
R
r
r
RR
Rr
Rr
rr
Evolutionary force #4: Genetic drift
Genetic drift – when allele frequencies changerandomly
Could be randomness of sexual reproduction, or any
other random event (natural disaster)
Smaller populations have larger changes potentially
Frequencies:
Before: R = 50%, r = 50% After: R = 23%, r = 77%
Genotype s # mice in desert # mice after
RR
0
0
Rr
4
7
Evolutionary force #4: Genetic drift
Bottleneck effect – natural disaster kills many
organisms, survivors
Founder effect – very small population
isolated from larger population in new area (ex: islands)
Genetic drift tends to reduce variety
So what causes evolution?
Let’s break the 5 assumptions one at a time
1.No gene flow (emigration or immigration)
2.No mutations occur
3.No natural selection is occurring
4.Population is large in number (infinitely large)
Evolutionary force #5: Sexual selection
Mate choice for particular characteristics
makes certain traits more prominent
Tends to be female choice of males in many
species
Evolutionary force #5: Sexual selection
S = sexy phenotype s = not so sexy
Genotypes # organisms before
# organisms after 10 generations
SS
3
14
Ss
10
12
ss
17
4
Before:
S = 26.7%
s = 73.3%
After:
Overall summary
Requirements for H-Wequilibrium
1) No gene flow (no
immigration / emigration) 2) No mutation
3) No natural selection 4) Infinitely large
population
5) Random mating
Evolutionary forces
1) Gene flow
2) Mutation
3) Natural selection 4) Genetic drift
Hardy-Weinberg equations
IF H-W equilibrium exists, we can calculate
allele frequencies and genotype frequencies in population if we know one component
Calculations always involve decimals, not
percentages
40% = 0.40 6.3% = 0.063
Equation #1:
p + q = 1 p = overall dominant allele % in gene pool q = overall recessive allele % in gene pool
Hardy-Weinberg equations
Organisms get alleles from mom and dad(2 homologous chromosomes)
(p + q) x (p + q) = 1 p = dominant allele q = recessive allele
Equation #2:
p2 + 2pq + q2 = 1
Gene pool idea
R = black
fur
r = sandy
fur
RR
Rr
Rr
Rr
rr
rr
rr
rr
rr
rr
rr
rr
rr
rr
rr
rr
rr
Rr
Rr
rr
(20 total
organisms)
14 / 20 = rr =
70%
7 / 40 R =
17.5%
1 / 20 = RR =
5%
5 / 20 = Rr =
25%
33 / 40 r =
82.5%
or p = 0.175
or q = 0.825
or p
2~
0.05
or 2pq ~
0.25
or q
2~
0.70
Gene pool
Hardy-Weinberg calculations
If we know one component of equation, we cansolve for other parts
Often, this component is the recessive phenotype –
q2
Ex: If 1 in 30,000 Americans has cystic fibrosis
(assume ~ 300,000,000 Americans), how many
Americans are carriers, assuming H-W equilibrium?
Cystic fibrosis H-W problem
Given q2, we can find q = √ q2
Then we can find p: p + q = 1
so p = 1 –q = 1 - √ q2
Then 2pq = 2(1 - √ q2) √ q2
2pq = 2 (1 – √0.00003)(√ 0.00003) = 0.0115
= 1.15%
1.15% of 300,000,000 Americans ~ 3,444,100
Cystic fibrosis H-W problem
Given q2, we can find q = √ q2 Then we can find p: p + q = 1
so p = 1 –q = 1 - √ q2
Then 2pq = 2(1 - √ q2) √ q2
2pq = 2 (1 – √0.00003)(√ 0.00003) = 0.0115 =
1.15%
1.15% of 300,000,000 Americans ~ 3,444,100
Solving without a calculator
Can you do these calculations?
2 x 0.60 x 0.40 = (0.30)2 =
(0.60)2 = √(0.04) =
Final reminder
These calculations assume population is in
H-W equilibrium