The evolution of populations

BIO 1510 Lecture 2

 On tuesday
• Concept of evolution proposed by Darwin:
Descent with modification
• Evidences previous to Darwin’s proposal:
extinctions (catastrophism), changes through
time (gradualism and fossil record), limited
survival within populations, and adaptation, and
biogographic variation (finches from Galapagos)
• Origin of the species: enviroment acting on
natural variations within populations-Natural
selection
• Support for evolution theory
 Today- Lecture 2
• How does evolution actually occurs?
• Genetic Variation Sourc
• The Hardy-Weiberg equilibrium and its
equation
• Populations and evolution
Natural Selection:
Genetic drift
Gene flow
Sexual selection
 The Smallest Unit of Evolution
• One misconception is that organisms
evolve during their lifetimes

• Natural selection acts on individuals, but

only populations evolve.
• A population is a group of individuals of
the same species living in the same
geographic area.
 Ground finches on Daphne Major Island
(Grant’s work)

During a drought,
large-beaked birds
were more likely to
crack large seeds
and survive
The finch population
evolved by natural
selection
Ground finches on Daphne Major
Island

During a drought,
large-beaked
birds were more
likely to crack
large seeds and
survive
The finch population
evolved by natural
selection

ENVIRONMENT (DROUGHT) BEAK SIZE = NATURAL SELECTION

 MICROEVOLUTION & MACROEVOLUTION

• Microevolution describes details on how

populations of organisms change over
generations.
– For polyploid organisms: It is is a change in allele
frequencies in a population over generations.

• Macroevolution describes patterns of changes

in groups of related species over periods of
geologic times (theousands to millions of years).
 Change over generations…. of
What?
• Phenotypic changes?....
• Gene changes?.......

• BOTH?…….
 GENETIC VARIATION
• Genetic variation is a pre-requisite for evolutionary
changes to occur.
• What type?:
– Mutations: one (Punctual) or multiple base
substitutions.
– Gene duplications- by error during Meiosis
– Deletions, insertions, translocation( movement of
gene onto another position).
– Sexual reproduction- gametes new gene
combiantions, random gametes encounter result in
“unique individual genetic combiantions”
• In real populations, allele and genotype
frequencies do change over time

• These changes can accumulate in time

resulting in evolutionary changes that
extended in generations can lead to the origin
of different species.
• In real populations, allele and genotype
frequencies do change over time
• These changes can accumulate in time
resulting in evolutionary changes that
extended in generations can lead to the origin
of different species.

HOW CAN WE TEST THAT A POPULATION

IS EVOLVING?
 The Hardy-Weinberg theorem
• Describes a non evolving
population.
• Used to test if a In a diploid organisms
Each pair of homologous chromosomes
population is evolving by Carry a copy of the same gene.
Each copy represents an alllele
estimating changes in
allele frecuencies p2  2pq  q2  1
• States that allele
P is the frecuency of allele A
frecuencies in the gene Q is the frecuency of allele B
pool of a population
remains constant unless
“agents” act upon them
Alelles, Loci, and genetic variation
 Gene pool
all copies of all
genes in all
members of a
population
 • The frequency of an allele in a population can
be calculated
– For diploid organisms, the total number of
alleles at a locus is the total number of
individuals times 2
– The total number of dominant alleles at a locus
is:
• 2 alleles for each homozygous dominant individual
• 1 allele for each heterozygous individual
(the same logic applies for recessive alleles)

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 • By convention, if there are 2 alleles at a locus,
p and q are used to represent their
frequencies
• The frequency of all alleles in a population will
add up to 1
– For example, p + q = 1

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 Alleles frequencies of the Parents
will determine allele frequencies of
the offspring
Calculating the allele frequencies of a particular loci
• For example, consider a population of
wildflowers that is incompletely dominant for
color:
– 320 red flowers (CRCR)
– 160 pink flowers (CRCW)
– 20 white flowers (CWCW)
• Calculate the number of copies of each allele:
– CR  (320  2)  160  800
– CW  (20  2)  160  200

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• The Hardy-Weinberg theorem describes a
hypothetical population that is not evolving

• If p and q represent the relative frequencies of

the only two possible alleles in a population at a
particular locus, then
– p2  2pq  q2  1
p2 and q2 represent the frequencies of the
homozygous genotypes
2pq represents the frequency of the heterozygous
genotype
 • Hardy-Weinberg equilibrium describes the
constant frequency of alleles in such a gene
pool
• Consider, for example, the same population
of 500 wildflowers and 100 alleles where
– p  freq CR  0.8
– q  freq CW  0.2

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 • To calculate the frequency of each allele:
– p  freq CR  800 / (800  200)  0.8
– q  freq CW  200 / (800  200)  0.2
• The sum of alleles is always 1
– 0.8  0.2  1

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 Figure 23.8
80% CR (p = 0.8) 20% CW (q = 0.2)

Sperm
CR (80%) CW (20%)

CR
(80%)
64% (p2) 16% (pq)
Eggs CRCR CRCW

CW 16% (qp) 4% (q2)

(20%) CRCW CWCW

64% CRCR, 32% CRCW, and 4% CWCW

Gametes of this generation:

64% CR R
+ 16% C R W = 80% CR = 0.8 = p
(from CRCR plants) (from C C plants)
4% CW W
+ 16% C R W = 20% CW = 0.2 = q
(from CWCW plants) (from C C plants)
Genotypes in the next generation:

64% CRCR, 32% CRCW, and 4% CWCW plants

 • If p and q represent the relative frequencies of
the only two possible alleles in a population at
a particular locus, then
– p2  2pq  q2  1
– where p2 and q2 represent the frequencies of
the homozygous genotypes and 2pq
represents the frequency of the heterozygous
genotype

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 Conditions for Hardy-Weinberg
Equilibrium
• The Hardy-Weinberg theorem describes a
hypothetical population that is not evolving
• In real populations, allele and genotype
frequencies do change over time

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 • The five conditions for nonevolving populations
are rarely met in nature:
1. No mutations
2. Random mating
3. No natural selection
4. Extremely large population size
5. No gene flow

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 • The five conditions for nonevolving populations
are rarely met in nature:

1. No mutations
2. Random mating
3. No natural selection Natural selection happens!!!

4. Extremely large population size Near extinctions

happens!!!
5. No gene flow

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 • Natural populations can evolve at some loci,
while being in Hardy-Weinberg equilibrium at
other loci

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 Applying the Hardy-Weinberg
Principle
• We can assume the locus that causes
phenylketonuria (PKU) is in Hardy-Weinberg
equilibrium given that:
1. The PKU gene mutation rate is low
2. Mate selection is random with respect to whether
or not an individual is a carrier for the PKU allele

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 3. Natural selection can only act on rare
homozygous individuals who do not follow
dietary restrictions
4. The population is large
5. Migration has no effect as many other
populations have similar allele frequencies

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 • The occurrence of PKU is 1 per 10,000 births
– q2  0.0001
– q  0.01
• The frequency of normal alleles is
– p  1 – q  1 – 0.01  0.99
• The frequency of carriers is
– 2pq  2  0.99  0.01  0.0198
– or approximately 2% of the U.S. population

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Three Mechanisms cause allele
frequency change:
• Natural Selection
• Genetic drift
• Gene flow
 Natural Selection
• Differential success in reproduction results
in certain alleles being passed to the next
generation in greater proportions
• For example, an allele that confers
resistance to DDT increased in frequency
after DDT was used widely in agriculture
 Directional, Disruptive, and
Stabilizing Selection
• Three modes of selection:
– Directional selection favors individuals at one
end of the phenotypic range
– Disruptive selection favors individuals at both
extremes of the phenotypic range
– Stabilizing selection favors intermediate
variants and acts against extreme phenotypes

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 Figure 23.13

Frequency of
individuals
Original population

Phenotypes (fur color)

Original Evolved
population population

(a) Directional selection (b) Disruptive selection (c) Stabilizing selection

 Natural Selection

• Evolution by natural selection involves

both change and “sorting”

New genetic variations arise by chance

Beneficial alleles are “sorted” and favored by
natural selection

• Only natural selection consistently

results in adaptive evolution
• Relative fitness is the contribution an
individual makes to the gene pool of the
next generation, relative to the
contributions of other individuals
• Selection favors certain genotypes by
acting on the phenotypes of certain
organisms
 The Founder Effect
• The founder effect occurs when a few
individuals become isolated from a larger
population
• Allele frequencies in the small founder
population can be different from those in the
larger parent population

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 The Bottleneck Effect
• The bottleneck effect is a sudden reduction in
population size due to a change in the
environment
• The resulting gene pool may no longer be
reflective of the original population’s gene pool
• If the population remains small, it may be further
affected by genetic drift

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 Figure 23.10-1

Original
population
 Figure 23.10-2

Original Bottlenecking
population event
 Figure 23.10-3

Original Bottlenecking Surviving

population event population
 • Understanding the bottleneck effect can
increase understanding of how human activity
affects other species

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 Case Study: Impact of Genetic Drift
on the Greater Prairie Chicken
• Loss of prairie habitat caused a severe
reduction in the population of greater prairie
chickens in Illinois
• The surviving birds had low levels of genetic
variation, and only 50% of their eggs hatched

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 Figure 23.11

Pre-bottleneck Post-bottleneck
(Illinois, 1820) (Illinois, 1993)

Greater prairie chicken

Range
of greater
prairie
(a)
chicken

Number Percentage
Location Population
of alleles of eggs
size
per locus hatched

Illinois
1930–1960s 1,000–25,000 5.2 93
1993 <50 3.7 <50

Kansas, 1998
750,000 5.8 99
(no bottleneck)

Nebraska, 1998 75,000–

5.8 96
(no bottleneck) 200,000

(b)
 Figure 23.11a

Pre-bottleneck Post-bottleneck
(Illinois, 1820) (Illinois, 1993)

Greater prairie chicken

Range
of greater
prairie
(a)
chicken
 Figure 23.11b

Number Percentage
Location Population
of alleles of eggs
size
per locus hatched

Illinois
1930–1960s 1,000–25,000 5.2 93
1993 <50 3.7 <50

Kansas, 1998
750,000 5.8 99
(no bottleneck)

Nebraska, 1998 75,000–

5.8 96
(no bottleneck) 200,000

(b)
 • Researchers used DNA from museum
specimens to compare genetic variation in the
population before and after the bottleneck
• The results showed a loss of alleles at several
loci
• Researchers introduced greater prairie
chickens from population in other states and
were successful in introducing new alleles and
increasing the egg hatch rate to 90%

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 Effects of Genetic Drift: A Summary
1. Genetic drift is significant in small populations
2. Genetic drift causes allele frequencies to
change at random
3. Genetic drift can lead to a loss of genetic
variation within populations
4. Genetic drift can cause harmful alleles to
become fixed

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 Gene Flow
• Gene flow consists of the movement of alleles
among populations
• Alleles can be transferred through the movement
of fertile individuals or gametes (for example,
pollen)
• Gene flow tends to reduce variation among
populations over time

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 • Gene flow can decrease the fitness of a population
• Consider, for example, the great tit (Parus major)
on the Dutch island of Vlieland
– Mating causes gene flow between the central and
eastern populations
– Immigration from the mainland introduces alleles
that decrease fitness
– Natural selection selects for alleles that increase
fitness
– Birds in the central region with high immigration
have a lower fitness; birds in the east with low
immigration have a higher fitness
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 Figure 23.12

Population in which the

60 surviving females Central
eventually bred population

50 Central NORTH SEA Eastern

Eastern population
Survival rate (%)

Vlieland,
40 the Netherlands
2 km
30

20

10

0
Females born Females born
in central in eastern Parus major
population population
 • Gene flow can increase the fitness of a population
• Consider, for example, the spread of alleles for
resistance to insecticides
– Insecticides have been used to target mosquitoes
that carry West Nile virus and malaria
– Alleles have evolved in some populations that
confer insecticide resistance to these mosquitoes
– The flow of insecticide resistance alleles into a
population can cause an increase in fitness

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THE END