Chapter 13

How Populations Evolve

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Clown, Fool, or Simply Well Adapted?

The blue-footed booby has many specialized characteristics that are very functional in water but less useful on land Such evolutionary adaptations are inherited traits that enhance an organism's ability to survive and reproduce in its particular environment Evolution is the changes in organisms over time
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First comes love Then comes baby!

DARWIN'S THEORY OF EVOLUTION 13.1 A sea voyage helped Darwin frame his theory of evolution
Pre-Darwinian ideas about the origin of species Early Greek philosophers: Simpler life forms preceded more complex ones Aristotle: Species are fixed and do not evolve; had a great impact on Western thinking Judeo-Christian biblical view: All species were individually designed by a divine creator
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 In the century prior to Darwin, only a few scientists questioned the belief that species are fixed

Buffon: The study of fossils suggested that Earth is older than 6,000 years, and fossil forms might be early versions of modern forms
Lamarck: Fossils are related to modern forms because life evolves; acquired characteristics are inherited

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Charles Darwin made a round-the-world sea voyage as a naturalist on HMS Beagle in the 1830s

Great Britain North America PACIFIC OCEAN ATLANTIC OCEAN

Europe

Asia

Africa
Equator

PACIFIC OCEAN

The Galpagos Islands

Pinta Marchena Santiago

PACIFIC OCEAN Genovesa Equator Daphne Islands

South America

Australia Cape of Good Hope Cape Horn Tierra del Fuego Tasmania New Zealand

Fernandina

Pinzn

Isabela

Santa Santa Cruz Fe Florenza

San Cristobal

0 0

40 km

Espaola

40 miles

 Darwin observed similarities between living and fossil organisms and the diversity of life on the Galpagos Islands

Darwin's experiences during the voyage helped him frame his ideas about evolution

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Frigate bird and giant tortoise

Marine iguana

Lyell's Principles of Geology led him to realize that still-operating natural forces gradually change Earth

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 After his return, Darwin began to document his observations and his new theory of evolution Alfred Wallace conceived a theory almost identical to Darwin's; both works were presented to the scientific community

Darwin's On the Origin of Species by Means of Natural Selection was published in 1859

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 "Descent with modification" summarizes Darwin's view of life All organisms are related through descent from a remote common ancestor Descendants spread into diverse habitats over millions of years and acquired adaptations to their environments The history of life resembles a tree with multiple branchings from a common trunk Species that are closely related share characteristics
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13.2 Darwin proposed natural selection as the mechanism of evolution

The essence of Darwin's theory of natural selection is differential success in reproduction Organisms produce more offspring than the environment can support Organisms vary in many characteristics that can be inherited Excessive numbers of organisms lead to a struggle for survival
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 Individuals whose characteristics are best adapted to their environment are more likely to survive and reproduce

The unequal ability of individuals to survive and reproduce leads to a gradual change in the characteristics of a population over generations
Natural selection is supported by evidence from artificial selection

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Controlled Breeding
Artificial selection is selective breeding to produce plants and animals that possess desirable traits Modern dogs descended from wolves In only a few thousand years, humans artificially selected for all breeds of modern dogs

LE 13-2c

African wild dog

Coyote

Wolf

Fox

Jackal

Thousands to millions of years of natural selection

Ancestral canine

Controlled Breeding
Humans have created tremendous variation in several species over relatively short periods of time through artificial selection Isnt it plausible that much larger changes could result from hundreds of millions of years of natural selection?

13.3 The study of fossils provides strong evidence for evolution

Fossils are the hard parts of organisms that remain after organic materials decay Rarely, an entire organism is fossilized

The fossil record strongly supports the theory of evolution

Changes in sea level and drying and refilling of lakes over time result in rock strata that trap organisms
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 Fossils appear in an ordered array within layers of sedimentary rocks The fossil record reveals that organisms have evolved in a historical sequence Many fossils link early extinct species with species living today

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13.4 A mass of other evidence reinforces the evolutionary view of life

Biogeography The geographic distribution of species suggested to Darwin that organisms evolve from common ancestors Isolated organisms resemble each other more than organisms in similar but distant places

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 Comparative anatomy Homologous structures are features that often have different functions but are structurally similar because of common ancestry

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LE 13-4a

Human

Cat

Whale

Bat

 Vestigial structures are remnants of structures that served important functions in an organism's ancestors

 Analogous structures are structures that are outwardly similar in appearance, but differ in their evolutionary origin

 Analogous structures result from convergent evolution rather than descent from a common ancestor Convergent evolution occurs when similar environmental pressures and natural selection give rise to similar (analogous) structures in distantly related organisms

 Comparative embryology Common embryonic structures in all vertebrates are evidence for common descent

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Lemur

pig

human

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Embryology
All vertebrate embryos possess genes that direct development of gill slits and a tail These genes were inherited from a common ancestor

Embryology
Adult fish retain gills and tail because the genes are active throughout their embryonic development Humans are born without gills and a tail because the genes are active only during early embryonic development

LE 13-4b

Pharyngeal pouches

Post-anal tail

Chick embryo

Human embryo

 Molecular biology Comparisons of DNA and amino acid sequences between different organisms reveal evolutionary relationships Molecular biology provides strong evidence that all life forms are related

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Molecular biology
All organisms share related biochemical processes:
All cells use DNA as genetic blueprint All use RNA, ribosomes, and approximately the same genetic code for translation All use roughly the same set of 20 amino acids to build proteins All use ATP to transfer energy

Molecular biology
Striking genetic similarities between organisms imply evolutionary relatedness e.g. the DNA nucleotide sequence of the human and mouse cytochrome c gene is very similar, suggesting shared ancestry

Cytochrome C gene-function in ETC

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CONNECTION 13.5 Scientists can observe natural selection in action

Examples of evolutionary adaptation observed over a short time Camouflage

Coloration in Trinidadian guppies

Pesticide resistance

Experimental introductions of Anolis sagrei lizards

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Camouflage

Coloration
Female guppies prefer to mate with brightly colored males; however, brightly colored males are more likely to be eaten by predators
Males found in areas lacking predators were brightly colored Males found in areas with predators were duller by comparison (predators eliminated brightly colored males before they could reproduce)

Coloration
Conclusion: When fewer predators are present, brighter coloration can evolve Conclusion was confirmed
Predators were introduced to previously predator-free areas (males were brightly colored) Within a few generations male guppies in those areas evolved to become less colorful

Pesticide Resistance
Numerous insect pests have evolved resistance to pesticides
Roaches developed resistance to Combat, an insecticide bait that acted as an agent of natural selection Resistant roaches possessed a rare mutation that caused them to dislike glucose, the main attractant in Combat

LE 13-5b

Chromosome with gene conferring resistance to pesticide

Pesticide application

Additional applications of the same pesticide will be less effective, and the frequency of resistant insects in the population will grow

Survivor

Pesticide Resistance
At least one insect species is resistant to every pesticide in existence

Experiments
Small groups of Anolis sagrei lizards were introduced onto 14 small Bahamian islands with thinly-branched bushes and no trees
Lizards were originally from Staniel Cay, an island with thickly-branched trees Their long legs were adaptive for maneuvering in these trees

The introduced lizards thrived and reproduced

Experiments
After 14 years, comparisons were made between lizards on the Bahamian islands and those of Staniel Cay Lizards on all 14 Bahamian islands had shorter, thinner legs

Experiments
Conclusion: Individuals with shorter, thinner legs evolved because they were able to escape predators better than their longer-legged ancestors in the new environment

 Examples of evolutionary adaptation reveal three key points about natural selection Natural selection is more of an editing process than a creative mechanism Natural selection is contingent on time and place Significant evolutionary change can occur in a short time

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POPULATION GENETICS AND THE MODERN SYNTHESIS

13.6 Populations are the units of evolution

Population
A group of individuals of the same species living in the same place at the same time May be isolated from other groups or concentrated

The smallest unit that can evolve

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 Population genetics Combines Darwin's and Mendel's ideas in studying how populations change genetically over time The modern synthesis
Connects population genetics with other sciences
Focuses on population as the unit of evolution and central role of natural selection
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 Studying evolution at the population level Evolution: change in the prevalence of certain heritable characteristics in a population over a span of generations Gene pool: the total collection of genes in a population at any one time Microevolution: a change in the relative frequencies of alleles in a gene pool Species: a group of populations capable of interbreeding and producing fertile offspring

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13.7 The gene pool of a nonevolving population remains constant over the generations
In a nonevolving population, the shuffling of alleles that accompanies sexual reproduction does not alter the genetic makeup of the population In Hardy-Weinberg equilibrium, the frequency of each allele in the gene pool will remain constant unless acted upon by other agents

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 For a population to be in Hardy-Weinberg equilibrium, it must satisfy five main conditions The population is very large The population is isolated Mutations do not alter the gene pool Mating is random All individuals are equal in reproductive success

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 The Hardy-Weinberg conditions are rarely met in nature We can follow alleles in a population to observe if Hardy-Weinberg equilibrium exists Hardy-Weinberg equilibrium provides a basis for understanding how populations evolve

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LE 13-7a

Webbing

No webbing

LE 13-7b

Phenotypes

Genotypes Number of animals (total = 500)

WW 320
320 500

Ww 160
160 500

ww
20
20 500

Genotype frequencies

= 0.64

= 0.32

= 0.04

Number of alleles in gene pool (total = 1,000)

640 W

160 W + 160 w

40 w

Allele frequencies

800 1,000

= 0.8 W

200 1,000

= 0.2 w

LE 13-7c Recombination of alleles from parent generation

Sperm W sperm p = 0.8 p2 W egg p = 0.8 WW = 0.64 w sperm q = 0.2 Ww pq = 0.16

Eggs w egg q = 0.2

wW qp = 0.16

q2

ww = 0.04

Next generation: Genotype frequencies Allele frequencies 0.64 WW 0.8 W 0.32 Ww 0.2 w 0.04 ww

CONNECTION 13.8 The Hardy-Weinberg equation is useful in public health science

Public health scientists use the HardyWeinberg equation to estimate frequencies of disease-causing alleles in the human population Example: phenylketonuria (PKU)

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13.9 In addition to natural selection, genetic drift and gene flow can contribute to evolution
Genetic drift: change in the gene pool of a population due to chance Can alter allele frequencies in a population The smaller the population, the greater the impact
Bottleneck effect: an event that drastically reduces population size Founder effect: colonization of a new location by a small number of individuals
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LE 13-9a

Original population

Bottlenecking event

Surviving population

 Gene flow: the movement of individuals or gametes between populations Can alter allele frequencies in a population Tends to reduce differences between populations

Natural selection
Best-adapted individuals have the most reproductive success Results in accumulation of traits that adapt a population to its environment
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CONNECTION 13.10 Endangered species often have reduced variation

Loss of genetic variability due to bottlenecking may reduce a population's ability to adapt to environmental change Particularly threatening to endangered species such as the cheetah

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VARIATION AND NATURAL SELECTION 13.11 Variation is extensive in most populations

Individual variation exists in all sexually reproducing populations Heritable variation results from a combination of genotype and environmental influences

Polymorphism: two or more forms of phenotypic characteristics

Geographic variation: variation of an inherited characteristic from place to place
May occur along a geographic continuum (a cline)
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13.12 Mutation and sexual recombination generate variation

Mutations-changes in the nucleotide sequence of DNA-can create new alleles Only mutations in cells that produce gametes can affect a population's gene pool A mutation may rarely improve adaptation to the environment and thus contribute to evolution Sexual recombination generates variation by shuffling alleles during meiosis
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LE 13-12a

Parents

A1

A1

A2

A3

Meiosis

A1 Gametes

A2

A3

LE 13-12b

A1
Gametes

A2

A3

Fertilization

Offspring, with new combinations of alleles

A1

A2
and

A1

A3

CONNECTION 13.13 The evolution of antibiotic resistance in bacteria is a serious public health concern
Natural selection has led to the evolution of antibiotic-resistant bacteria Overuse and misuse of antibiotics has contributed to the proliferation of antibioticresistant strains Example: tuberculosis

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13.14 Diploidy and balancing selection preserve variation

Diploidy (two sets of chromosomes) helps to prevent populations from becoming genetically uniform Recessive alleles are "hidden" from natural selection and remain in the population

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 Balancing selection allows two or more phenotypic forms in a population Balanced polymorphism may result from
Heterozygote advantage; example: sicklecell disease

Frequency-dependent selection

Neutral variation provides no apparent advantage or disadvantage Example: fingerprints

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13.15 The perpetuation of genes defines evolutionary fitness

Evolutionary fitness is the relative contribution an individual makes to the gene pool of the next generation Survival of genes depends on production of fertile offspring Selection indirectly adapts a population to its environment by acting on phenotype

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13.16 Natural selection can alter variation in a population in three ways

Stabilizing selection: favors intermediate phenotypes Directional selection: acts against individuals at one of the phenotypic extremes Disruptive selection: favors individuals at both extremes of the phenotypic range

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LE 13-16
Frequency of individuals Original population

Phenotypes (fur color) Original population Evolved population

Stabilizing selection

Directional selection

Disruptive selection

13.17 Sexual selection may produce sexual dimorphism

Sexual dimorphism The distinction in appearance between males and females of a species

Sexual selection
The determining of "who mates with whom"

Leads to the evolution of secondary sexual characteristics that may give individuals an advantage in mating
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13.18 Natural selection cannot fashion perfect organisms

There are at least four reasons why natural selection cannot produce perfection Organisms are limited by historical constraints Adaptations are often compromises Chance and natural selection interact Selection can only edit existing variations
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