1. The Evolution of Populations
Chapter 23
The Evolution of Populations

2. Overview: The Smallest Unit of Evolution
One misconception is that organisms evolve, in the Darwinian sense, during their lifetimes
Natural selection acts on individuals, but only populations evolve
Genetic variations in populations contribute to evolution
Microevolution is a change in allele frequencies in a population over generations

3. Is this finch evolving by natural selection?
Fig. 23-1
Is this finch evolving by natural selection?
Figure 23.1 Is this finch evolving by natural selection?

4. Concept 23.1: Mutation and sexual reproduction produce the genetic variation that makes evolution possible
Two processes, mutation and sexual reproduction, produce the variation in gene pools that contributes to differences among individuals

5. Genetic Variation
Variation in individual genotype leads to variation in individual phenotype
Not all phenotypic variation is heritable
Natural selection can only act on variation with a genetic component

6. Diet related appearences
Fig. 23-2
Diet related appearences
(a)
(b)
Feed on the flowers
Feed on the leaves
Figure 23.2 Nonheritable variation

7. Variation Within a Population
Both discrete and quantitative characters contribute to variation within a population
Discrete characters can be classified on an either-or basis (colors of pea flowers)
Quantitative characters vary along a continuum within a population (hair color)

8. Population geneticists measure polymorphisms in a population by determining the amount of heterozygosity at the gene and molecular levels
Average heterozygosity measures the average percent of loci that are heterozygous in a population
Nucleotide variability is measured by comparing the DNA sequences of pairs of individuals

9. Variation Between Populations
Fig. 23-3
Variation Between Populations
Most species exhibit geographic variation
*fused chromosomes haven't caused a genetic change
*therefore the mice look the same
Island of Madeira
8.11 1
2.4
3.14
5.18 6
7.13
8.11
9.12
10.16
13.17 19 XX
Figure 23.3 Geographic variation in isolated mouse populations on Madeira
Mus musculus 1
2.19
3.8
4.16
5.14
6.7
9.10
11.12
13.17
15.18 XX

10. Cline is a graded change in a trait along a geographic axis 1.0 0.8
Fig. 23-4
1.0
0.8
0.6
Cline is a graded
change in a trait
along a geographic
axis
Ldh-B b allele frequency
0.4
0.2
Figure 23.4 A cline determined by temperature 46 44 42 40 38 36 34 32 30
Latitude (°N)
Maine
Cold (6°C)
Georgia
Warm (21°C)

11. Mutation
Mutations are changes in the nucleotide sequence of DNA
Mutations cause new genes and alleles to arise
Only mutations in cells that produce gametes can be passed to offspring

12. A point mutation is a change in one base in a gene
Point Mutations
A point mutation is a change in one base in a gene
The effects of point mutations can vary:
Mutations that result in a change in protein production are often harmful
Mutations that result in a change in protein production can sometimes increase the fit between organism and environment

13. Mutations That Alter Gene Number or Sequence
Chromosomal mutations that delete, disrupt, or rearrange many loci are typically harmful
Duplication of large chromosome segments is usually harmful
Duplication of small pieces of DNA is sometimes less harmful and increases the genome size
Duplicated genes can take on new functions by further mutation

14. Mutation Rates
Mutation rates are low in animals and plants
The average is about one mutation in every 100,000 genes per generation
Mutations rates are often lower in prokaryotes and higher in viruses

15. Sexual Reproduction
Sexual reproduction can shuffle existing alleles into new combinations (crossing over)
In organisms that reproduce sexually, recombination of alleles is more important than mutation in producing the genetic differences that make adaptation possible

16. Concept 23.2: The Hardy-Weinberg equation can be used to test whether a population is evolving
A population is a localized group of individuals capable of interbreeding and producing fertile offspring
A gene pool consists of all the alleles for all loci in a population
A locus is fixed if all individuals in a population are homozygous for the same allele

17. Shared area Porcupine herd Porcupine herd range Fortymile herd range
Fig. 23-5
Porcupine herd
MAP
AREA
CANADA
ALASKA
Beaufort Sea
NORTHWEST
TERRITORIES
Porcupine
herd range
Shared area
Figure 23.5 One species, two populations
Fortymile
herd range
ALASKA
YUKON
Fortymile herd

18. 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 x 2
The total number of dominant alleles at a locus is 2 alleles for each homozygous dominant individual plus 1 allele for each heterozygous individual; the same logic applies for recessive alleles

19. The frequency of all alleles in a population will add up to 1
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

20. The Hardy-Weinberg Principle
The Hardy-Weinberg principle describes a population that is not evolving
If a population does not meet the criteria of the Hardy-Weinberg principle, it can be concluded that the population is evolving

21. Hardy-Weinberg Equilibrium
The Hardy-Weinberg principle states that frequencies of alleles and genotypes in a population remain constant from generation to generation
In a given population where gametes contribute to the next generation randomly, allele frequencies will not change
Mendelian inheritance preserves genetic variation in a population

22. Alleles in the population Frequencies of alleles
Fig. 23-6
Alleles in the population
Frequencies of alleles
Gametes produced
p = frequency of
Each egg:
Each sperm:
CR allele = 0.8
q = frequency of
80%
chance
20%
chance
80%
chance
20%
chance
CW allele = 0.2
Figure 23.6 Selecting alleles at random from a gene pool

23. Hardy-Weinberg equilibrium describes the constant frequency of alleles in such a gene pool
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

24. Figure 23.7 The Hardy-Weinberg principle
80% CR ( p = 0.8)
20% CW (q = 0.2)
Sperm CR
(80%) CW
(20%) CR
(80%)
Eggs
64% ( p2)
CR CR
16% ( pq)
CR CW
4% (q2)
CW CW CW
(20%)
16% (qp)
CR CW
64% CR CR, 32% CR CW, and 4% CW CW
Figure 23.7 The Hardy-Weinberg principle
Gametes of this generation:
64% CR    +     16% CR    =   80% CR  = 0.8 = p
4% CW      +    16% CW    =  20% CW = 0.2 = q
Genotypes in the next generation:
64% CR CR, 32% CR CW, and 4% CW CW plants

25. Conditions for Hardy-Weinberg Equilibrium
The Hardy-Weinberg theorem describes a hypothetical population, not always occurs in nature
In real populations, allele and genotype frequencies do change over time

26. The five conditions for nonevolving populations are rarely met in nature:
No mutations
Random mating
No natural selection
Extremely large population size
No gene flow

27. Applying the Hardy-Weinberg Principle
Natural populations can evolve at some loci, while being in Hardy-Weinberg equilibrium at other loci
We can assume the locus that causes phenylketonuria (PKU) is in Hardy-Weinberg equilibrium given that:
The PKU gene mutation rate is low
Mate selection is random with respect to whether or not an individual is a carrier for the PKU allele

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

29. The occurrence of PKU is 1 per 10,000 births
q2 =
q = 0.01
The frequency of normal alleles is
p = 1 – q = 1 – 0.01 = 0.99
The frequency of carriers is
2pq = 2 x 0.99 x 0.01 =
or approximately 2% of the U.S. population

30. Concept 23.3: Natural selection, genetic drift, and gene flow can alter allele frequencies in a population
Three major factors alter allele frequencies and bring about most evolutionary change:
Natural selection
Genetic drift
Gene flow

31. Natural Selection
Differential success in reproduction results in certain alleles being passed to the next generation in greater proportions

32. Genetic Drift
The smaller a sample, the greater the chance of deviation from a predicted result (Tay-Sachs)
Genetic drift describes how allele frequencies fluctuate unpredictably from one generation to the next
Genetic drift tends to reduce genetic variation through losses of alleles

33. Generation 1 Generation 2 Generation 3 p (frequency of CR) = 0.7
Fig
CR CR
CR CR
CW CW
CR CR
CR CR
CR CW
CR CW
CR CR
CR CR
CW CW
CR CR
CR CR
CW CW
CR CR
CR CR
CR CW
CR CW
CR CR
CR CR
CR CR
CR CR
CR CW
CW CW
CR CR
Figure 23.8 Genetic drift
CR CR
CR CR
CR CR
CR CW
CR CW
CR CW
Generation 1
Generation 2
Generation 3
p (frequency of CR) = 0.7
p = 0.5
p = 1.0
q (frequency of CW ) = 0.3
q = 0.5
q = 0.0

34. 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

35. 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
Gene flow is a change in the allele frequency due to transfer of alleles into or out of the pool

36. Example of Bottleneck effect: Attwater Chicken
From 1,000,000 in 1900s to less
than 100 today
Lack of grassland, environmental
pressure (predators, fireants)
Low genetic pool

37. Fig
Gene flow
Figure Gene flow and human evolution

38. Gene flow can decrease the fitness of a population
In bent grass, alleles for copper tolerance are beneficial in populations near copper mines, but harmful to populations in other soils
Windblown pollen moves these alleles between populations
The movement of unfavorable alleles into a population results in a decrease in fit between organism and environment

39. Gene flow can increase the fitness of a population
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

40. Concept 23.4: Natural selection is the only mechanism that consistently causes adaptive evolution
Natural selection brings about adaptive evolution by acting on an organism’s phenotype
Fitness
The phrases “struggle for existence” and “survival of the fittest” are misleading as they imply direct competition among individuals
Reproductive success is generally more subtle and depends on many factors

41. 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

42. 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

43. Frequency of individuals
Fig
Original population
Frequency of individuals
Phenotypes (fur color)
Original
population
Evolved
population
Figure Modes of selection
(a) Directional selection
(b) Disruptive selection
(c) Stabilizing selection

44. The Key Role of Natural Selection in Adaptive Evolution
Natural selection increases the frequencies of alleles that enhance survival and reproduction
Adaptive evolution occurs as the match between an organism and its environment increases

45. predators and surprise prey
Fig
Enables it to hide from
predators and surprise prey
(a) Color-changing ability in cuttlefish
Movable bones
Figure Examples of adaptations
Allows them to swallow food
much larger than the mouth
(b) Movable jaw
bones in
snakes

46. Because the environment can change, adaptive evolution is a continuous process
Genetic drift and gene flow do not consistently lead to adaptive evolution as they can increase or decrease the match between an organism and its environment

47. Sexual Selection
Sexual selection is natural selection for mating success
It can result in sexual dimorphism, marked differences between the sexes in secondary sexual characteristics

48. Intrasexual selection is competition among individuals of one sex (often males) for mates of the opposite sex
Intersexual selection, often called mate choice, occurs when individuals of one sex (usually females) are choosy in selecting their mates
Male showiness due to mate choice can increase a male’s chances of attracting a female, while decreasing his chances of survival

49. The Preservation of Genetic Variation
Various mechanisms help to preserve genetic variation in a population
Diploidy maintains genetic variation in the form of hidden recessive alleles

50. Heterozygote advantage occurs when heterozygotes have a higher fitness than do both homozygotes
Natural selection will tend to maintain two or more alleles at that locus
The sickle-cell allele causes mutations in hemoglobin but also confers malaria resistance

51. Plasmodium falciparum (a parasitic unicellular eukaryote) 7.5–10.0%
Fig
Frequencies of the
sickle-cell allele
0–2.5%
Figure Mapping malaria and the sickle-cell allele
2.5–5.0%
5.0–7.5%
Distribution of
malaria caused by
Plasmodium falciparum
(a parasitic unicellular eukaryote)
7.5–10.0%
10.0–12.5%
>12.5%

52. Frequency-Dependent Selection
In frequency-dependent selection, the fitness of a phenotype declines if it becomes too common in the population
Selection can favor whichever phenotype is less common in a population
Moth changing color due to pollution

53. Neutral Variation
Neutral variation is genetic variation that appears to confer no selective advantage or disadvantage
For example,
Variation in noncoding regions of DNA
Variation in proteins that have little effect on protein function or reproductive fitness

54. Why Natural Selection Cannot Fashion Perfect Organisms
Selection can act only on existing variations, favors the fittest
Evolution is limited by historical constraints, does not "occur" out of the blue
Adaptations are often compromises
Chance, natural selection, and the environment interact
The End