Biol 1115

Chapter 22: Evolution by Natural Selection

Mário Moniz de Sá, PhD

Chapter 22 Opening Roadmap
Learning Objectives: You should be able to …

• Define evolution, fitness, and adaptation using the biological definitions.

• Describe the nature of the evidence regarding (1) whether species change
through time and (2) whether they are related by common ancestry.

• Assess whether Darwin’s four postulates are true in any given example,
explain to a friend why evolution must occur if all four are true, and explain
whether evolution will occur if any of the four are not true.

• Identify common misconceptions about evolution and adaptation, and give

examples to illustrate why they are not true. (For example: Is evolution
progressive? Are all traits adaptive? Does evolution result in perfection? Does
evolution proceed by the accumulation of changes occurring by chance? Do
organisms develop traits that they need in their environment?)
 Introduction to Evolution by Natural Selection
• Evolution by natural selection is one of the best-supported and most
important theories in modern biology
– One of the great ideas in science
– One of the five key attributes of life:
§ Populations of organisms evolve, or change through time

• Charles Darwin published On the Origin of Species by Means of

Natural Selection in 1859
– Radically different from the idea of special creation, which held
that
1. Species are independent (unrelated)
2. Life on Earth is young (~ 6000 years old)
3. Species are immutable (incapable of change)

• Scientific theory
1. Is not a guess
2. Explains a broad class of observations and is widely supported
by evidence

• Scientific theories are often comprised of two components:

1. Pattern—observations about natural world
2. Process—mechanism that produces that pattern
 The Evolution of Evolutionary Thought
• Theory of evolution by natural selection is often described as revolutionary

• A scientific revolution replaces an existing idea about nature with a radically

different idea
– The idea of special creation dominated Western thought about the nature
of organisms for over 2000 years

• The Greek philosopher Plato

claimed that
– Every organism was an
example of a perfect
essence, or type, created by
God
– Types were essentially
unchanging

• Today, philosophers and biologists

refer to ideas like this as
typological thinking
 Aristotle and the Scale of Nature
• Aristotle ordered known organisms into the linear
great chain of being (or scale of nature), in which
– Species were fixed types
– Species were organized into a sequence
based on increasing size and complexity
– Sequence started with minerals and lower
plants
– Humans were at the top of the chain

• In the 1700s Aristotle’s ideas were still popular.

The central claims were that
– Species were fixed types as Plato had
proposed
– Some species are higher—in the sense of
being more complex or “better”—than others
Lamarck and the Idea of Evolution
as Change Through Time
• In 1809 Jean-Baptiste de Lamarck was the first to
propose a formal theory of evolution:
– Based on Aristotle’s great chain of being, simple
organisms originate at the base of the chain by
spontaneous generation
– Organisms evolve by moving up the chain over time
– Lamarckian evolution is progressive—producing
larger, more complex or “better” species over time

• Lamarck suggested that the process responsible for this

pattern was the inheritance of acquired characters
– As individuals develop, their phenotype changes in
response to environmental challenges
– phenotypic changes are then passed on to their
offspring
– Example: Giraffes develop long necks from
stretching to reach food and produce offspring with
long necks
 Darwin and Wallace and Evolution by Natural
Selection
• Darwin and Wallace proposed that change in species through time
– Does not follow a linear, progressive pattern
– Is based on variation among individuals in populations
§ A population consists of individuals of the same species living in
the same area at the same time
– Individuals with certain traits produce more offspring than others
without these traits

• This view uses population thinking rather than typological thinking

– Darwin claimed that variation among individuals in a population was
the key to understanding evolution

• The theory of evolution by natural selection was revolutionary for several

reasons:
1. It overturned the idea that species are static and unchanging
2. It replaced typological thinking with population thinking
3. It was scientific
§ It proposed a mechanism that could account for change
through time
§ Its predictions could be tested through observation and
experimentation
The Pattern of Evolution: Have Species
Changed, and Are They Related?

• Darwin described evolution as

descent with modification
– Change over time produced
modern, modified species
from ancestral species
• The pattern component of the
theory of evolution by natural
selection predicts that
1. Species change through time
2. Species are related by
common ancestry
 Evidence for Change Through
Time
• Fossils are traces of organisms that
– lived in the past

• The many fossils that have been found and described

in the scientific literature
– make up the fossil record

• Fossils were initially organized according to their

relative ages

• Sedimentary rocks
– Form from sand or mud
– Form in layers
§ Younger layers are deposited on top of older
layers

• Scientists created a geologic time scale based on

fossils’ relative positions in layers of sedimentary rock
– Geologic time is divided into eons, eras, periods,
and epochs

• The geologic record indicated that Earth was much,

much older than 6000 years
The Vastness of Geologic Time
 The Vastness of Geologic Time
• Researchers can now assign absolute ages to the
relative ages in the geologic time scale by using
radioactive decay
– The steady rate of conversion of unstable
“parent” atoms into stable “daughter” atoms

• Radiometric dating is based on

– Observed decay rates
– Ratio of parent to daughter atoms in newly
formed rocks
– Ratio of parent to daughter atoms in a particular
rock sample

• Radiometric dating data indicate that

– Earth is about 4.6 billion years old
– The earliest signs of life are found in rocks about
3.4–3.8 billion years old
 Extinction Changes the
Species Present over Time
• Many fossils provide evidence for extinct
species
– species that no longer exist
• Darwin interpreted extinction as evidence
that
– Species are dynamic
– The array of species living on Earth
has changed through time
• Recent analyses of the fossil record
suggest that
– Over 99% of all the species that
have ever lived are now extinct
– Extinctions have occurred
continuously throughout Earth’s
history
Transitional Features Link Older and Younger Species
• Early scientists observed that fossil species are strikingly similar to
living species in the same geographic areas
– This pattern became known as the “law of succession”

• Darwin interpreted this pattern as evidence that

– Species changed through time
– Extinct and living forms were related and represented
ancestors and descendants

• Transitional features are traits in a fossil species that are

intermediate between ancestral and derived species
– These transitional forms provide strong evidence for change
through time
§ For example, fossils show a gradual change from the
aquatic fin to the terrestrial limb

• Data like these are consistent with predictions from the theory of
evolution
– If the traits observed in more recent species evolved from
traits in more ancient species, then transitional forms are
expected to occur
 Vestigial Traits are Evidence of
Change Through Time
• A vestigial trait in an organism is
– A reduced or incompletely developed
structure that has no (or reduced) function
but clearly similar to functioning organs or
structures in closely related species

• Examples of vestigial traits include

– Nonfunctional hip and leg bones in some
snakes and whales
– Reduced wings in flightless birds
– Eye sockets in eyeless cave-dwelling fish
– Brief eggshell formation and nonfunctional
“egg tooth” in some marsupial species
– Coccyx bone (vestigial tail) and goose
bumps in humans

• Vestigial traits are

– Inconsistent with the idea of special
creation
– Evidence that the characteristics of species
have changed over time
Current Examples of Change
Through Time
• Hundreds of contemporary populations have been
documented undergoing evolutionary changes.
For example:
– Bacteria have evolved resistance to drugs
– Insects have evolved resistance to pesticides
– Weedy plants have evolved resistance to
herbicides
– Bird migration, insect emergence, and
blooming of flowering plants have evolved in
response to climate change

• Species continue to change.

• Data from the fossil record and contemporary

species refute the hypothesis that species are
immutable

• There is also mounting contemporary scientific

evidence of the relatedness of species by
common ancestry
 Similar Species are Found in the Same Geographic Area

• There are often striking similarities among island species

– For example, mockingbirds that Darwin collected from the Galápagos
Islands were superficially similar, but different islands had distinct species

• Darwin proposed that the mockingbirds were similar because they had
descended from a common ancestor

• Recent DNA analyses support Darwin’s hypothesis

 Similar Species are Found in the Same Geographic Area
• A phylogenetic tree is a diagram that illustrates the ancestor–descendant
relationships among taxa

• Researchers compared mockingbird DNA and placed the birds on a

phylogenetic tree, which shows that
– The different Galápagos mockingbird species are each others’ closest
living relatives
– They share a single common ancestor
Homology is Evidence of Descent from a Common Ancestor
• Another line of evidence comes from
homologies

• Homology is a similarity that exists in species

descended from a common ancestor

• Homology can be recognized and studied at

three interacting levels:
1. Genetic
2. Developmental
3. Structural

• Genetic homology is a similarity in the DNA

nucleotide sequences, RNA nucleotide
sequences, or amino acid sequences
– For example, the eyeless gene in fruit
flies and the Aniridia gene in humans are
so similar that their resulting amino acid
sequences are 90% identical
 Homology Is Evidence of Descent from a Common Ancestor

• The three levels of homology interact

– Genetic homologies cause the developmental
homologies observed in embryos
– Chemicals that cause cancer in humans can
often be identified by testing their effects on
mutation rates in bacteria, yeast, zebrafish,
mice, and other model organisms
– Drugs intended for human use can be tested on
mice or rabbits if the proteins or other gene
products targeted by the drugs are homologous

• Hypotheses about homology can be tested

experimentally. For example:
– A fruit fly embryo received a mouse gene that
signals where eyes should form
– A fruit fly eye formed in the location where the
mouse gene was expressed
 Table 22.2 Evidence for Evolution
Prediction 1: Species Are Not Static, but Change through Time
• Life on Earth is ancient. The fossil record shows how life has changed over time.
• Fossil (extinct) species frequently resemble living species found in the same area.
• Transitional features document change in traits through time.
• Vestigial traits are common.
• The characteristics of populations vary within species and can be observed
changing today.

Prediction 2: Species Are Related, Not Independent

• Similar species often live in the same geographic area.
• Homologous traits are common and are recognized at three levels:
1. genetic (gene structure and the genetic code)
2. developmental (embryonic structures and processes)
3. structural (morphological traits in adults)
• The formation of new species from pre-existing species can be observed today.
 Evolution’s “Internal Consistency”—The Importance of
Independent Data Sets
• Internal consistency is
– The observation that data from independent sources agree in supporting
predictions made by a theory
– Perhaps the most powerful evidence for any scientific theory, including
evolution by natural selection

• Evidence for the evolution of the cetaceans—whales and dolphins—illustrates

the idea of internal consistency
– Fossil cetaceans are identified by unique ear bones
– A phylogeny of cetaceans indicates a gradual transition between
terrestrial and aquatic forms

• Relative and absolute dating both support the order of species indicated in the
phylogeny

• DNA comparisons indicate that hippos are the closest living relative of
cetaceans and that they share a semiaquatic ancestor

• Vestigial hip and limb bones are found in some adult whales and dolphin
embryos
 Data on Evolution from Independent Sources Are Consistent
• Data from many different sources are much more consistent with evolution than with special
creation
• Descent with modification is a more successful and powerful scientific theory
• It explains observations—such as vestigial traits and the close relationships among
species on neighbouring islands—that special creation does not
The Process of Evolution: How Does Natural
Selection Work?
• While many researchers had proposed evolution
as a pattern in nature long before Darwin,

• Darwin’s contribution was describing a process,

– Called natural selection,
– That could explain the pattern of descent with
modification

• Darwin based his idea of natural selection on

– Observations exploring and documenting the
diversity of plants and animals around the
globe.
– Work of geologist Charles Lyell and,
– The work of Thomas Robert Malthus.

• Artificial selection in pigeon breeding

– Darwin crossbred pigeons and observed how
characteristics were passed on to offspring
– He concluded that the diverse pigeon breeds
all descended from wild pigeons
 Darwin’s Inspiration
• Thomas Robert Malthus wrote the book An Essay on the
Principle of Population
– Described a “struggle for existence”
§ Many more individuals are born than can survive
§ Thus, people compete for resources

• Darwin combined several ideas to arrive at his concept of

natural selection
– His observations of artificial selection
– The notion of struggle for existence in natural populations
– Variation in natural populations

• Both Darwin and Wallace arrived at the same idea, but Darwin’s
name is widely associated with the concept of natural selection
– He thought of it first
– He provided extensive evidence in On the Origin of Species
 Darwin’s Four Postulates
• Darwin broke the process of evolution by natural selection into four criteria, or
postulates:
1. Individuals in a population vary in their traits
2. Some of these differences are heritable; they are passed on to offspring
3. In each generation, many more offspring are produced than can survive
§ Only some will survive long enough to reproduce
§ Some will produce more offspring than others
4. Individuals with certain heritable traits are more likely to survive and
reproduce
§ Natural selection occurs when individuals with certain traits produce
more offspring than do individuals without those traits
§ The individuals are selected naturally, by the environment

• The selected traits will increase in frequency in the population from one generation
to the next, causing evolution
– Evolution is a change in the allele frequencies of a population over time

• Evolution is thus a logical outcome of the four postulates

• Modern biologists condense Darwin’s four steps into two statements: Evolution by
natural selection occurs when
1. Heritable variation leads to
2. Differential reproductive success
The Biological Definitions
of Fitness and Adaptation

• Biological fitness
– The ability of an individual to
produce surviving, fertile offspring
relative to that ability in other
individuals in the population

• Adaptation
– A heritable trait that increases an
individual’s fitness in a particular
environment relative to individuals
lacking that trait

• Selection
– Differential reproduction as a
result of heritable variation
 Evolution in Action: Recent Research
on Natural Selection
• The theory of evolution by natural selection is testable
– Document heritable variation and
– Differential reproductive success

• Examples:
– Drug resistance in bacteria
– Beak size and shape and body size changes in the Galápagos finches
 Case Study 1: How Did Mycobacterium Tuberculosis
Become Resistant to Antibiotics?

• The bacterium Mycobacterium tuberculosis causes

tuberculosis (TB)
– TB infects the lungs, often causing death
– TB was once as great a public health issue as
cancer is now

• TB receded in importance in the early 1900s due to

– Improved nutrition
– The development of antibiotics

• However
– In the late 1980s, rates of TB surged
– In 1993 the World Health Organization declared
TB a global health emergency
– Strains of the bacteria responsible for the
increase were resistant to the once-effective
antibiotics
 A Patient History
• A young man was diagnosed with an active TB infection
– He was given several antibiotics for 6 weeks and doses
of the antibiotic rifampin for an additional 33 weeks
§ Tests showed the infection was cleared after 10
months
– Two months later, he was readmitted to the hospital
with a recurrence of the symptoms
§ He died 10 days later, despite additional antibiotic
treatment

• Tests showed that the M. tuberculosis bacteria in his lungs

were completely resistant to rifampin

• DNA from the rifampin-resistant bacteria was found to have a

single point mutation in a gene called rpoB
– rpoB codes for a component of RNA polymerase
– The mutation caused a change in amino acid sequence
resulting in a change in enzyme shape

• Rifampin works by binding with the M. tuberculosis RNA

polymerase and interfering with transcription
– The mutation prevents rifampin from binding
Alleles That
Confer Drug
Resistance
Increase in
Frequency
When Drugs
Are Used
 Testing Darwin’s Postulates
• Variation existed in the population
– Due to mutation, both resistant and nonresistant strains of TB were present before
administration of the drug

• The variation was heritable

– The variation in the phenotypes of the two strains was due to variation in their genotypes

• There was variation in reproductive success

– Only a tiny fraction of M. tuberculosis cells survived the first round of antibiotics long enough to
reproduce

• Selection occurred
– The cells with the drug-resistant allele had higher reproductive success

• This example shows how natural selection acts on individuals because

individuals experience differential success

• It also shows how only populations evolve, as allele frequencies change in

populations, not individuals

• Understanding evolution by natural selection requires population thinking

 Drug Resistance: A Widespread Problem

• Resistance to a wide variety of

insecticides, fungicides, antibiotics,
antiviral drugs, and herbicides has
evolved in hundreds of insects, fungi,
bacteria, viruses, and plants

• In every case, evolution has occurred

because individuals with the heritable
ability to resist some chemical
compound were present in the original
population
– As the susceptible individuals die
from the pesticide, herbicide, or
drug, the resistance alleles
increase in frequency
 Canadian Issues: Evolution in Action: Do
Hunting and Fishing Select for Undesirable
Traits?
• Atlantic cod (Gadus morhua)
population declined 99% between
1962 and 1992
– estimated two billion cod were
lost
– Atlantic cod with a genetic
predisposition to mature at larger
size or greater age were likely to
be caught before they were able
to reproduce.
– Cod that matured early, at a
small size, had greater
reproductive success.

• Strong directional selection for early-

maturing, small adult cod.
Canadian Issues 22.1: Evolution in Action: Do
Hunting and Fishing Select for Undesirable
Traits?
• Small adult cod suffer a higher natural mortality, have a shorter life span, and
produce fewer offspring than the large cod that were targeted by fishers

• Unlike natural predators, humans harvest other predators;

– Humans also target large age and size classes.
– Natural predators focus on “the newly born and the nearly dead,”
– Humans prefer to take large, healthy reproductive adults.
– Humans also take a far higher proportion of the harvested populations

• Humans cause far more rapid phenotypic changes than other predators do.

• Human-harvested systems examined in the study showed phenotypic changes

that were 300 percent higher than those in natural systems.

• Humans are causing large and rapid changes in the size and life history of
harvested populations.

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 Case Study 2: Why Do Beak Sizes and Shapes
Vary in Galápagos Finches?
• Peter and Rosemary Grant
have conducted long-term
research on the population of
medium ground finches found
on Isle Daphne Major of the
Galápagos Islands

• They found that beak form and

body size are heritable in these
birds
Selection During Drought Conditions (1 of 2)
• During the Grants’ research, a major drought led 84% of the ground finch
population to die of starvation

• The research team realized that the die-off was a natural experiment
– Natural experiments allow researchers to compare treatment groups
created by an unplanned change in conditions
– The Grants could test whether natural selection occurred by comparing
the population before and after the drought

• In only one generation, natural selection led to a measurable change in the

characteristics of the population
– The finch population’s average beak depth had increased
§ Deeper beaks were adaptive due to food availability during the
drought
§ Large, deep beaks were an adaptation for cracking large fruits and
seeds
– Alleles that led to the development of deep beaks had increased in
frequency
Figure 22.14 A Natural
Experiment: Changes
in a Medium Ground
Finch Population in
Response to a
Change in the
Environment (a
Drought)

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 Continued Changes in the Environment, Continued
Selection, Continued Evolution
• In 1983
– Over 7 months more than 10
times the average annual rainfall
occurred
– altered relative plant abundance
– Small individuals with small,
pointed beaks had exceptionally
high reproductive success
§ Alleles associated with small,
pointed beaks increased in
frequency

• Grants have documented continued

evolution in response to continued
changes in the environment
 Which Genes Are Under Selection?
• Many characteristics, including beak size,
are polygenic—many genes each exert a
relatively small effect. For example:
– Beak depth is a polygenic trait
– Research has shown that the amount
of expression of a gene called Bmp4
correlates with beak depth
– Thus, this gene is thought to be
under selection in the medium ground
finch

• This illustrates a connection between

natural selection on phenotype and
evolutionary change in genotype

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Table 22.3 Common Misconceptions, Corrected
• Perhaps the most important point to clarify about natural selection is that during the
process, individuals do not change—only the population does. For example:
– During the drought, individual beaks did not change
§ The average beak depth increased over time because deep-beaked
individuals had greater reproductive success
– Individual M. tuberculosis cells did not change
§ The frequency of the rifampin-resistant allele increased in the population over
time
Table 22.3 Common Misconceptions, Corrected
• Evolution by natural selection is not goal directed. Mutations do not occur to solve
problems—mutations are random.
– Due to errors during DNA synthesis.

• Adaptations do not occur because organisms want or need them

Table 22.3 Common Misconceptions, Corrected
• Evolution is not progressive—meaning it does not produce “better” or more
complex organisms

• In fact, complex traits are routinely lost or simplified over time as a result of
evolution by natural selection.
– Populations that become parasitic are particularly prone to loss of
complex traits.
Acclimatization Is Not Adaptation
• Acclimatization
– Occurs when an individual’s phenotype changes in
response to changes in the environment
– The individual’s genotype remains fixed
– The changes are not passed on to offspring, because
no alleles have changed

• In contrast, adaptation occurs when the allele

frequencies in a population change in response to
natural selection
Figure 22.17 Evolution Produces a Tree of Life, Not a
Progressive Ladder of Life

• Scientifically, there is no such thing as

“higher” or “lower” organisms

• Mosses may be a more ancient group

than flowering plants, but neither group is
higher or lower than the other.
– Different suite of adaptations.
Traits Are Not Always Adaptive
• Vestigial traits confer no known benefit
– Organisms possessing them do not have higher fitness
than those without
• Silent mutations
– Changes in the DNA sequence that do not result in a
change in the amino acid sequence of the protein
encoded by the gene
– Extremely common
– Do not change the phenotype and thus cannot be
acted on by natural selection and are not adaptive
 Traits Are Genetically Constrained

• Selection is not able to optimize all aspects of a trait

due to certain genetic constraints
• Genetic correlation occurs
– When selection favouring alleles for one trait causes a
correlated but suboptimal change in an allele for another
trait
– Because of pleiotropy, in which a single allele affects
multiple traits

• Lack of genetic variation can also constrain evolution,

because natural selection can work only on existing
variation in a population
 Fitness Trade-Offs Exist

• A fitness trade-off is a compromise between traits, in

terms of how those traits perform in the environment
• Because selection acts on many traits at once, every
adaptation is a compromise,
• Examples of fitness trade-offs include compromises
between
– The size of eggs or seeds that an individual makes and the
number of offspring it can produce
– Rapid growth and long life span
– Bright colouration and tendency to attract predators
Traits Are Historically Constrained

• Because all traits evolve from previously existing

traits,
– adaptations are constrained by history
• Natural selection acts on structures that originally
had a very different function
– the best solution possible given an important historical
constraint
Traits Are Environmentally Constrained

• Natural selection often occurs in the context of a

changing environment
– Abiotic and biotic features can fluctuate
§ Over time
§ Over the geographic range of the population

• Not all traits are adaptive, and even adaptive traits

are constrained by genetic and historical factors.
– natural selection is not the only process that causes
evolutionary change
 The End
Where’s the python?

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