11.

2 Natural Selection in Populations

Key Concept Populations, not individuals, evolve.
MAIN IDEAS
VOCABULARY Natural selection acts on distributions of traits.
normal distribution Natural selection can change the distribution of a trait in one of three ways.
microevolution
directional selection
stabilizing selection Connect to Your World
disruptive selection How do you describe a person’s appearance? Perhaps you use height, hair color, and
eye color. These traits are often used in descriptions because these traits vary widely
among humans. In this section, you will learn about the way natural selection can act
on such variation.

MAIN IDEA
Natural selection acts on distributions of traits.
Any time you stand in a large crowd of people, you are likely to observe a wide
CONNECT TO range of heights. Imagine organizing this crowd across a football field accord-
Genetics ing to each individual’s height, with very short people at one end, people of
As you learned in the chapter
average height in the middle, and very tall people at the other end. You would
Extending Mendelian Genetics, soon notice a pattern in distribution of the human height trait. Relatively few
single-gene traits are expressed in people would be at each extreme height, very short or very tall. A majority of
either one distinct form or people of medium height would be in the middle.
another. However, the range of
phenotypes common for most This type of distribution, in which the frequency is highest near the
traits is the result of polygenic mean value and decreases toward each extreme end of the range, is called a
traits, which are controlled by
multiple genes. normal distribution. When these frequency values are graphed, the result is a
bell-shaped curve like the one you see in Figure 2.1.
For some traits, all phenotypes provide an equal chance of
Figure 2.1 normal distribution survival. The distribution for these traits generally shows a normal
mean distribution. Phenotypes near the middle of the range tend to be
most common, while the extremes are less common. However,
environmental conditions can change, and a certain phenotype
may become an advantage. Natural selection favors individuals with
Frequency

this phenotype. These individuals are able to survive and reproduce

at higher rates than individuals with less favorable phenotypes.
Therefore, alleles associated with favorable phenotypes increase
in frequency through differential reproductive success.
Synthesize What other types of data might follow a normal
Range of variable distribution?

318 Unit 4: Evolution

 MAIN IDEA
Natural selection can change the distribution of
a trait in one of three ways.
Microevolution is the observable change in the allele frequencies of a popula-
tion over time. Microevolution occurs on a small scale—within a single
population. One process that can lead to microevolution is natural selection.
Natural selection can change the distribution of a trait along one of three
paths: directional, stabilizing, or disruptive selection. Such changes can have
major effects on how a population looks and behaves.

Directional Selection
A type of selection that favors phenotypes at one extreme of a trait’s range CONNECT TO
is called directional selection. Directional selection causes a shift in a popula- Bacteria
tion’s phenotypic distribution. An extreme phenotype that was once rare in Although many bacteria are
a population becomes more common. As shown in Figure 2.2, during direc- helpful to other organisms, some
tional selection, the mean value of a trait shifts in the direction of the more do cause disease. You will learn
more about how bacteria can
advantageous phenotype. evolve and become resistant
The rise of drug-resistant bacteria provides a classic example of this type to antibiotics in the chapter
of selection. Before antibiotics were developed in the 1940s, a trait for varying Viruses and Prokaryotes.
levels of drug resistance existed among bacteria. At the time, there was no The rise of drug-resistant bacteria is a
advantage to having drug resistance. But once antibiotics came into use, the prime example of natural selection.
resistant bacteria had a great advantage. Before antibiotics, some bacteria had
varying levels of drug resistance, but
The early success of antibiotics in controlling infectious diseases led to it wasn't advantageous. However,
overuse of these drugs. This overuse favored even more resistant phenotypes. with the widespread use of antibiotics,
New drugs were then developed to fight the resistant bacteria. This resulted in drug-resistant bacteria gained an
the evolution of “superbugs” that are highly resistant to many drugs. Today, edge. Overuse of antibiotics led to
even more resistant strains, resulting
over 200 types of bacteria show some degree of antibiotic resistance. in "superbugs" highly resistant to
many drugs. Today, over 200 bacteria
types display antibiotic resistance.
FIGURE 2.2 Directional Selection
Directional selection occurs when one extreme phenotype is favored
by natural selection.
mean mean Today, scientists continue
to research new drugs devel-
oped to treat infection-causing
Original distribution
bacteria such as Enterococcus
faecalis, which is resistant to
Frequency

Antibiotic drugs put many antibiotics.

pressure on bacteria
populations.
©Mark Thomas/Photo Researchers, Inc.

Distribution after
directional selection

Low drug resistance High drug resistance

Chapter 11: The Evolution of Populations 319

 Stabilizing Selection
The gall fly and its predators provide an excellent example of stabilizing
selection. During stabilizing selection, the intermediate phenotype is favored
and becomes more common in the population. That is, the distribution
becomes stable at the intermediate phenotype rather than shifting toward one
of the extremes. In the case of gall flies, something in nature selects against
gall
phenotypes at both extremes of the trait’s range.
Gall flies lay their eggs in developing shoots of the tall goldenrod plant.
The fly larvae produce a chemical that causes the plant tissue to swell around
them. Figure 2.3 shows the resulting mass of plant tissue, called a gall. The gall
serves as a home where the larvae can develop. There is a range of phenotypes
for body size in gall-fly larvae. Each body size causes a certain size gall to
form, and each of the two main predators of gall flies specializes on a specific
gall size.
• Downy woodpeckers attack larger galls and feed on the larvae inside.
FIGURE 2.3 The gall fly and
the goldenrod plant have a
• The parasitic wasp lays its own eggs inside small galls. After the wasp
parasitic relationship. The fly larvae emerge from the eggs, they eat the gall-fly larvae.
benefits by receiving shelter
and food during its larval stage, In this situation, selective pressure from predators works against fly

(tl) ©Warren G. Abrahamson, Ph.D; (inset) ©Peter Harris, Agriculture and Agri-Food Canada/Forestry Images; (c) ©M.F. Claridge; (br) ©Tom Reichner/Shutterstock
while the goldenrod is harmed, phenotypes that produce galls at both extremes, large and small. As a result,
growing more slowly than a flies that produce middle-sized galls become more common. As you can see in
gall-free goldenrod.
figure 2.4, over time, stabilizing selection results in a higher frequency of flies
Gall flies lay eggs in tall goldenrod shoots, and their
larvae cause the plant to form galls, serving as protective
that produce middle-sized galls.
homes. Predators, like downy woodpeckers, attack
larger galls, while parasitic wasps target smaller ones. Stabilizing selection increases the number of individuals with intermediate
This results in selective pressure against extreme gall
sizes. Stabilizing selection favors gall flies producing
phenotypes. Notice, however, that selection against both extremes decreases
middle-sized galls, leading to an increase in their the genetic diversity of the gall-fly population. Flies that produce small and
frequency. However, it reduces genetic diversity by
making small and large gall-producing flies less large galls become less common. In some populations, these
common, and they may even disappear in some
populations.
extreme phenotypes may be lost altogether.

FIGURE 2.4 Stabilizing Selection

Stabilizing selection occurs when intermediate phenotypes are favored
by natural selection.
narrow & higher
mean

Original distribution
wasp woodpecker
Woodpeckers and wasps
put pressure on gall-fly
Frequency

populations.
Distribution after
stabilizing selection

Small gall size Large gall size

320 Unit 4: Evolution

 selective agent: factors in the environment

Disruptive Selection FIGURE 2.5 Disruptive Selection

Disruptive selection occurs when both extreme
Disruptive selection occurs when both extreme
phenotypes are favored, while individuals with phenotypes are favored by selection.
intermediate phenotypes are selected against
by something in nature. As you can see in
figure 2.5, the middle of the distribution is
disrupted. One example of this type of selection
involves feather color in male lazuli buntings, a

Frequency
bird species native to North America.
Young male lazuli buntings vary widely in the
brightness of their feathers, ranging from dull
brown to bright blue. Dominant adult males are
those with the brightest blue feathers on their
heads and backs. These birds have their pick of Brown Blue
the best territories. They also are most successful
at attracting females. However, for young bun- Original distribution
tings, the brightest blue and dullest brown males
Dominant adult males
are more likely to win mates than males with put pressure on young
bluish brown feathers. males in the bunting
population.
Research suggests that dominant adult males
are aggressive toward young buntings that they Distribution after
see as a threat, including bright blue and bluish disruptive selection
brown males. The dullest brown birds can there-
Adult male lazuli bunting
fore win a mate because the adult males leave
them alone. Meanwhile, the bright blue birds In figure 2.5, disruptive selection disrupts the middle
attract mates simply because of their color. of a distribution. This phenomenon is observed in
male lazuli buntings, where young males vary
Both extreme phenotypes are favored in this situation, while intermediate widely in feather color, ranging from dull brown to
bright blue. Dominant adult males with the brightest
forms are selected against. The bluish brown males are not as well adapted to blue feathers have advantages in securing
©C. Allan Morgan/Peter Arnold, Inc.

territories and attracting females. However, for

compete for mates because they are too blue to be left alone by adult males, but young buntings, the extremes—bright blue and
not blue enough to win a mate based on color alone. By favoring both extreme dullest brown males—have better chances of
mating, while bluish brown males face aggression
phenotypes, disruptive selection can lead to the formation of new species. from dominant adults. This leads to disruptive
selection favoring extreme phenotypes and, in some
cases, can contribute to the formation of new
Apply If bluish brown coloring became advantageous for young males, what type species.
of selection would likely occur in a lazuli bunting population?
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11.2 Formative Assessment CONNECT TO

Genetics
Reviewing  Main Ideas Critical thinking 5. For polygenic traits, a smooth
1. In terms of phenotypes, describe 3. Analyze How might the extinction curve results when the range
what is meant by the phrase of downy woodpeckers affect of phenotypes is plotted
“distribution of traits.” the phenotypic distribution within a against frequency. If you
population of gall flies? were to plot the frequencies
2. What are the three ways in which
4. Analyze How might overfishing of two phenotypes of a
natural selection can change a
of large pink salmon select for smaller single-gene trait, you would
distribution of traits?
body size in subsequent generations? end up with a double bar
graph. Explain why.

Chapter 11: The Evolution of Populations 321