1 Evolution is an ongoing process. 2 Darwin journeyed to a new

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1 Evolution is an ongoing process.
8.1 We can see evolution occurring right before our eyes.
2 Darwin journeyed to a new idea.
8.2 Before Darwin, most people believed that all species
had been created separately and were unchanging.
8.3 A job on a ’round-the-world survey ship allowed
Darwin to indulge and advance his love of nature.
8.4 Observing geographic similarities and differences
among fossils and living plants and animals, Darwin
developed a theory of evolution.
8.5 In 1858, after decades of mulling and procrastinating,
Darwin published his thoughts on natural selection.
3 Four mechanisms can give rise to evolution.
8.6 Evolution occurs when the allele frequencies in a
population change.
8.7 Mutation—a direct change in the DNA of an
individual—is the ultimate source of all genetic variation.
8.8 Genetic drift is a random change in allele frequencies
in a population.
8.9 Migration into or out of a population may change
allele frequencies.
8.10 When three simple conditions are satisfied, evolution
by natural selection is occurring.
4 Through natural selection, populations of organisms
can become adapted to their environment.
8.11 Traits causing some individuals to have more offspring
than others become more prevalent in the population.
8.12 Organisms in a population can become better
matched to their environment through natural selection.
8.13 Natural selection does not lead to perfect organisms.
8.14 Artificial selection is a special case of natural selection.
8.15 Natural selection can change the traits in a population
in several ways.
8.16 Natural selection can cause the evolution of complex
traits and behaviors.
5 The evidence for the occurrence of evolution is
overwhelming.
8.17 The fossil record documents the process of natural
selection.
8.18 Geographic patterns of species distributions reflect
their evolutionary histories.
8.19 Comparative anatomy and embryology reveal
common evolutionary origins.
8.20 Molecular biology reveals that common genetic
sequences link all life forms.
8.21 Laboratory and field experiments enable us to watch
evolution in progress.
8
Evolution
and Natural
Selection
Darwin’s
dangerous idea
1 Evolution is an
ongoing process.
8 • 1---------------------------------------------------- We can see evolution occurring right before our eyes.
What’s the longest that you’ve ever gone without food?
Twenty-four hours? Thirty-six hours? If you’ve ever gone
hungry that long, you probably felt as if you were going to die
of starvation, but humans can survive days, even weeks,
without food. In 1981, for example, 27-year-old Bobby Sands,
a political prisoner in Northern Ireland, went on a hunger
strike. Forsaking all food and consuming only water, he
gradually deteriorated and ultimately died—after 66 days
without food.
Each hour without food is even more dangerous when you’re
tiny. How long do you think a fruit fly can last without food?
The answer is just under a day—20 hours, give or take a few
(Figure 8-1). Fruit flies can’t live long without food because
their tiny bodies don’t hold very large caloric reserves. But
could you breed fruit flies that could live longer than 20 hours
on average? Yes.
First, start with a population of 5,000 fruit flies. In biology, the
term “population” has a specific meaning: a population is a
group of organisms of the same species living in a particular
geographic region. The region can be a small area like a test
tube or a large area like a lake. In your experiment, your
population occupies a cage in your laboratory.
From these 5,000 fruit flies, you choose only the “best” fruit
flies to start the next generation, with “best” meaning the 20%
of flies that can survive the longest without food. Choosing
these long-lived, hungry flies can be accomplished in three
simple steps:
1. Remove the food from the cage with the population of
5,000 flies.
2. Wait until 80% of the flies starve to death and then put a
container of food into the cage.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
population be even higher? In fact, it would. With each new
generation, you would see a slight increase, and with the fifth
generation, the population’s average survival would be about
32 hours.
One benefit to conducting research on fruit flies is that they
have very short lives—they reach maturity at about two weeks
of age and live for one month, on average. An experiment
such as this one can be continued for many generations. After
60 generations of allowing the flies that are “best” at surviving
without food to reproduce, how has the population changed?
Amazingly, the average fly in the resulting population can
survive for more than 160 hours without food (Figure 8-2). In
60 generations, the flies have gone from a starvation resistance
of less than a day to one of nearly a week! At the point where
the food is removed, the flies in generation 60 are noticeably
fatter than the flies in generation 0 were.
Figure 8-1 Starvation resistance in fruit flies.
3. After the surviving flies eat, they’ll have the energy to
reproduce. When they do, collect the eggs they lay and
transfer those eggs to a new cage.
You can now start a new generation populated only by the
offspring of those fruit flies able to survive without food for
the longest amount of time. When these eggs hatch, do you
think the flies in this new generation will live longer than 20 hours without food? When this experiment is done, the
flies do indeed show increased resistance to starvation. The
average fly in the new generation can live for about 23 hours
without food. And again, some of the new flies survive for
more than 23 hours, some for less. The surviving flies tend to
be those that pack on the fat when the food is available. The
new starvation-resistance curve for this second generation is
similar to that of the original flies, but shifted to the right a bit
so the highest part of the curve occurs at 23 hours, indicating
that most flies of the new generation survive for an average of
23 hours without food.
What if you kept repeating these three steps? Start each new
generation using eggs only from the fruit flies in the top 20%
of starvation resistance. After five generations, would the
average starvation-resistance time for a fly from your
What happened? In a word: evolution. That is, there was a
genetic change in the population of fruit flies living in the
cage. Every fly in the generation 60 population, even the fly
with the worst starvation resistance, is still more than seven
times better at resisting starvation than the best fly in the
original population. This evolution is the result of natural
selection. We’ll discuss natural selection in more detail later
in this chapter, but, in short, it is the consequence of certain
individual organisms in a population being born with
characteristics that enable them to survive better and
reproduce more than the offspring of other individuals in the
population. In this experiment, the 20% of fruit flies that were
the most starvation resistant had a huge reproductive
advantage over less-resistant flies because they were the only
flies in the population that survived to reproduce.
This experiment answers a question that is sometimes
perceived as complex or controversial: does evolution occur?
The answer is an unambiguous yes. We can watch it happen in
the lab whenever we want. Recall from our discussion of the
scientific method (see Chapter 1) that for an experiment’s
results to be valid, they must be reproducible. To be certain of
these results, researchers carried out the starvation-resistance
experiment described above five separate times. The results
were the same every time.
In this experiment we changed starvation resistance, but what
if we used dogs instead of flies and, instead of allowing the
most starvation-resistant individuals to reproduce, we allowed
only the smallest dogs to reproduce? Would the average body
size of individual dogs decrease over time? Yes. What if the
experiment were done on rabbits and only the fastest rabbits
escaped death from predation by foxes? Would the average
running speed in rabbits increase over time? Yes. We know
Evidence for Evolution
Figure 8-2 Evolution in action: increasing fruit fly resistance to starvation.
these answers because each of these experiments has already
been done.
In this chapter we examine how evolution can occur and the
type of changes it can cause in a population. We also review
the five primary lines of evidence that indicate that evolution
and natural selection are processes that help us to clarify all
other ideas and facts in biology. Let’s begin our investigation
with a look at how the idea of evolution by natural selection
was developed. This knowledge will help us better understand
why this idea is regarded as one of the most important ideas in
human history, why it is sometimes considered a dangerous
idea, and why it generates emotional debate.
TAKE-HOME MESSAGE
8 1
•
The characteristics of individuals in a population can change
over time. We can observe such change in nature and can
even cause such change to occur.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
2 Darwin
journeyed to a
new idea.
8 • 2--------------------------------------------------- Before Darwin, most people believed that all species had
been created separately and were unchanging.
Charles Darwin grew up in an orderly world. When it came
to humans and our place in the world, in the mid-19th
century the beliefs of nearly everyone were virtually
unchanged from the beliefs of people who had lived more
than 2,000 years earlier. Biblical explanations were sufficient
for most natural phenomena: the earth was thought to be
about 6,000 years old. With the occasional exception of a
flood or earthquake or volcanic eruption, the earth was
believed to be mostly unchanging. People recognized that
organisms existed in groups called species. (In Chapter 10 we
discuss in more detail what a species is; for now, we’ll just say
that individual organisms in a given species can interbreed
with each other but not with members of another species.)
People also believed that all species, including humans, had
been created at the same time and that, once created, they
never changed and never died out. This was pretty much what
Aristotle had believed more than two millennia earlier.
Before he left this world, Darwin had thrown into question
long and dearly held beliefs about the natural world and
forever changed our perspective on the origins of humans and
our relationship to all other species. He didn’t smash the
worldview to pieces all at once, though, and he didn’t do it by
himself (Figure 8-3).
In the 1700s and 1800s, scientific thought was advancing at a
rapid pace. In 1778, the respected French naturalist Georges
Buffon began to shake things up by suggesting that the earth
must be about 75,000 years old. He arrived at this age by
estimating that 75,000 years was the minimum time required
for the planet to cool from a molten state. In the 1790s,
Georges Cuvier began to explore the bot­­toms of coal and
slate mines and found fossil remains that had no obvious
similarity to any living species. The implications of Cuvier’s
discoveries were unthinkable, since biblical accounts did not
allow for species to be wiped out. Cuvier’s publications
documented giant fossils (including the Irish elk, the
mastodon, and the giant ground sloth) that bore no
resemblance to any currently living animals (Figure 8-4).
These fossils allowed only one explanation: extinction was a
fact. Troubling as this obser­vation was for the prevailing
worldview, it was only the beginning.
Evidence for Evolution
Figure 8-3 Scientists who shaped Darwin’s thinking.
Not only was it starting to seem that species
could disappear from the face of the earth, but
the biologist Jean-Baptiste Lamarck suggested
in the early 1800s that living species might
change over time. Although his ideas about the
mechanisms by which this change might occur
were wrong—he thought that change came
about through the use or disuse of features—his
willingness to question previously sacred “truths” contributed
to an atmosphere of unfettered scientific thought in which it
was possible to challenge convention.
Perhaps the heretical ideas that most inspired Darwin were
those of the geologist Charles Lyell. In his 1830 book,
Principles of Geology, Lyell argued that geological forces had
shaped the earth and were continuing to do so, producing
mountains and valleys, cliffs and canyons, through gradual but
relentless change. This idea that the physical features of the
earth were constantly changing would most closely parallel
Darwin’s idea that the living species of the earth, too, were
gradually—but constantly—changing.
TAKE-HOME MESSAGE
8 2
•
In the 18th and 19th centuries, scientists began to overturn
the commonly held beliefs that the earth was only about
6,000 years old and that all species had been created
separately and were unchanging. These gradual changes in
scientists’ beliefs helped shape Charles Darwin’s thinking.
Figure 8-4 Extinction occurs. Deep
in coal mines, Cuvier discovered the
fossilized remains of very large animals
no longer found on earth.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
8 • 3--------------------------------------------------- A job on a ‘round-the-world survey ship allowed Darwin
to indulge and advance his love of nature.
Charles Darwin was born into a wealthy family in England in
1809 and his early life was unremarkable. Charles was never at
the top of his class and professed to hate schoolwork.
Nonetheless, at 16, he went to the University of Edinburgh to
study medicine, following his father’s steps. He was bored,
though, and finally left at the end of his second year, when the
prospect of watching gruesome surgeries (in the days before
anesthesia) was more than he could bear.
Upon the abrupt ending of Charles’s medical career, his father
urged him to pursue the ministry and sent him to study
theology at Cambridge University. Although he never felt
great inspiration in his theology studies, Darwin was in heaven
at Cambridge, where he could pursue his real love, the study
of nature. While at Cambridge, he spent most of his time
reading nature books, collecting specimens, and developing
close relationships with many of his professors.
FPO
Figure 8-6 Charles
Darwin. Darwin found his life’s
calling in exploration and study
of the natural world.
Shortly after graduation in 1831, Darwin—with significant
help from his botany professor—landed his dream job, a
position as a “gentleman companion” for the captain of the
HMS Beagle. The Beagle, a 90-foot sailing vessel, was on a
five-year, ’round-the-world surveying expedition (Figure 8-5).
This job came as a huge relief to the young Darwin, who was
not really interested in becoming a minister.
Once on the Beagle, Darwin found that he didn’t actually like
sea travel. He was seasick for much of his time on board,
where he shared a room that was much tinier than a dorm
room. He wrote to his cousin: “I hate every wave of the
ocean with a fervor which you . . . can never understand.” To avoid nausea, Charles spent as much time as possible on
shore. It was there that he found his true calling (Figure 8-6).
Darwin loved fieldwork. At each stop that the Beagle made, he
would eagerly investigate the new worlds he found. In Brazil,
he was enthralled by tropical forests. In Patagonia, near the
southernmost part of South America, he explored beaches and
cliffs, finding spectacular fossils from huge extinct mammals.
Elsewhere he explored coral reefs and barnacles, always
packing up specimens to bring back to museums in England
and recording his observations for later use. He was like a
schoolboy on permanent summer vacation.
TAKE-HOME MESSAGE
Figure 8-5 “Like a schoolboy on permanent summer vacation.”
In 1831, Darwin set out on the HMS Beagle on a five-year surveying
expedition that took him around the globe.
8 3
•
After initially training in medicine and theology, Charles
Darwin was able to focus on studying the natural world
when, in 1831, he got a job on a ship conducting a five-year,
‘round-the-world survey.
Evidence for Evolution
8 • 4--------------------------------------------------- Observing geographic similarities and differences among
fossils and living plants and animals, Darwin developed a
theory of evolution.
The only book Darwin brought with him on the Beagle was
Lyell’s Principles of Geology, which he read again and again and
discussed with anyone who would listen. Intrigued by the
book’s premise that the earth has been, and continues to be,
constantly changing, Darwin’s mind was ripe for fresh ideas—
particularly as he began observing seemingly inexplicable
things. How could he explain finding marine fossils high in
the Andes Mountains, hundreds of miles from the nearest
ocean? The idea that the earth was an ever-changing planet
was seeping into Darwin’s mind. That idea would serve him
well when, a few years into its journey, the Beagle stopped at
the Galápagos Islands, off the northwest coast of South
America.
This group of volcanic islands was home to many unusual
species, from giant tortoises to extremely docile lizards that
made so little effort to run away that Darwin had to avoid
stepping on them. Darwin was particularly intrigued by the
wide variety of birds, especially the finches, which seemed
dramatically more variable than those he had seen in other
locations.
Darwin noticed two important and unexpected patterns on
his voyage that would be central to his discovery of a
mechanism for evolution. The first involved the finches he
collected and donated to the Zoological Society of London.
Darwin had assumed that they were the equivalent of tall and
short, curly-haired and straight-haired people. That is, he
thought that all the finches were of the same species but with
different physical characteristics or traits, such as body size,
beak shape, or feather color. The staff of the Zoological
Society, however, could see from the birds’ physical differences
that there were 13 unique species—a different species from
every one of the Galápagos Islands that Darwin had visited.
Figure 8-7 Darwin observed unexpected patterns.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
Moreover, although the birds were different species, they all
resembled very closely the single species of finch living on the
closest mainland, in Ecuador (Figure 8-7).
This resemblance seemed a suspicious coincidence to Darwin.
Perhaps the island finches resembled the mainland species
because they used to be part of the same mainland population.
Over time they may have separated and diverged from the
original population and gradually formed new—but similar—
species. Darwin’s logic was reasonable, but his idea flew in the
face of all of the scientific thinking of the day.
The second important but unexpected pattern Darwin noted
was that, throughout his voyage, at every location there was a
striking similarity between the fossils of extinct species and
the living species in that same area. In Argentina, for instance,
he found some giant fossils from a group of organisms called
“glyptodonts.” The extinct glyptodonts looked just like
armadillos—a species that still flourished in the same area. Or
rather, they looked like armadillos on steroids: the average
armadillo today is about the size of a house cat and weighs 10 pounds, while the glyptodonts were giants. At 10 feet long
and more than 4,000 pounds, they were larger than an
automobile (see Figure 8-7).
If glyptodonts had lived in South America in the past, and
armadillos were currently living there, why was only one of
the species still alive? And why were the glyptodont fossils
found only in the same places that modern armadillos lived?
Darwin deduced that glyptodonts resembled armadillos
because they were their ancestors. Again, it was a logical
deduction, but it contradicted the scientific dogma of the day
that species were unchanging and extinction did not occur.
The wheels were turning in Darwin’s head, but he didn’t have
a great “Eureka!” moment until several years after his voyage,
when he was back in England. He was reading “for
amusement” a book called Essay on the Principle of Population
by the economist Thomas Malthus. Malthus prophesied doom
and gloom for populations—including humans—based on his
belief that populations had the potential to grow much faster
than food supplies could. Darwin speculated that, rather than
the future holding certain catastrophe for all, maybe the best
individuals would “win” in the ensuing struggle for existence
and the worst would “lose.” If so, he suddenly saw that
“favourable variations would tend to be preserved, and
unfavourable ones to be destroyed.” Everything was falling
into place for Darwin, who wrote: “Here then I had at last
got a theory by which to work.”
In 1842, Darwin finally hammered out a first draft of his ideas
in a 35-page paper, written in pencil, and fleshed it out over
the next couple of years. He knew that his idea was
important—so important that he wrote a letter to his wife
instructing her, in the case of his sudden death (an odd
request, given that he was only 35 years old), to give his
“sketch” to a competent person, with about $1,000 and all of
his books, so that this person could complete it for him.
He also had an inkling that his ideas would rock the world. In
a letter to a close friend, he wrote: “At last gleams of light
have come, and I am almost convinced (quite contrary to the
opinion I started with) that species are not (it is like confessing
to murder) immutable.” But “confessing to murder” was
apparently more than Darwin was ready for. Inexplicably, he
put his sketch into a drawer, where it remained for 14 years.
TAKE-HOME MESSAGE
8 4
•
Darwin noted unexpected patterns among fossils he
discovered and living organisms he observed while on
the voyage of the Beagle. Fossils resembled but were
not identical to the living organisms in the same area in
which they were found. And finch species on each of the
Galápagos Islands differed from each other in small but
significant ways. These observations helped Darwin to
develop his theory of how species might change over time.
Evidence for Evolution
8 • 5--------------------------------------------------- In 1858, after decades of mulling and procrastinating,
Darwin published his thoughts on natural selection.
It came in the form of a letter from a young British biologist
in the throes of malaria-induced mania in Malaysia. In the
letter, Alfred Russel Wallace, who had also read Malthus’s book
on population growth, laid out a clear description of the
process of evolution by natural selection (Figure 8-8). He
asked Darwin to “publish it if you think it is worthy.” Crushed
at having been scooped, Darwin wrote that “all my originality
will be smashed.” He had numerous friends among the most
prominent scientists of the day, however, and they arranged for
a joint presentation of Wallace’s and Darwin’s work to the
Linnaean Society of London. As a result, both Darwin and
Wallace are credited for the first description of evolution by
natural selection.
Darwin then sprang into action, rapidly putting together his
thoughts and observations and completing, 16 months later,
a full book. In 1859, he published Origin of Species (its full
title is On the Origin of Species by Means of Natural Selection,
or the Preservation of Favoured Races in the Struggle for Life)
(Figure 8-9).
The book was an instant hit, selling out on its first day,
provoking public discussion and debate and ultimately causing
a wholesale change in the scientific understanding of
Figure 8-9 Reconsidering the
world. Commonly held ideas before
and after Darwin.
evolution. Where once the worldview was of a young earth,
populated by unchanging species all created at one time, with
no additions or extinctions, now there was a new dynamic
view of life on earth: species could and did change over time,
and as some species split into new species, others became
extinct.
Darwin’s theory has proved to be among the most important
and enduring contributions in all of science. It has stimulated
an unprecedented diversity of theoretical and applied research
programs and it has withstood repeated experimental and
observational testing. With the background of Darwin’s
elegant idea in hand, we can now examine its details.
TAKE-HOME MESSAGE
Figure 8-8 Alfred Russel Wallace. Darwin and Wallace independently
identified the process of evolution by natural selection.
10 8 5
•
After putting off publishing his thoughts on natural
selection for more than 15 years, Darwin did so only after
Alfred Russel Wallace independently came up with the same
idea. The two men published a joint presentation on their
ideas in 1858, and Darwin published a much more detailed
treatment in Origin of Species in 1859, sparking wide debate
and discussion of natural selection.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
3 Four
mechanisms
can give rise to
evolution.
8 • 6--------------------------------------------------- Evolution occurs when the allele frequencies in a
population change.
Suppose you were put in charge of a large population of tigers
in a zoo.Virtually all of them are orange and brown with
black stripes. Occasionally, though, unusual all-white tigers are
born. This white phenotype is the result of the possession of a
rare pair of alleles that suppresses the production of most fur
pigment (Figure 8-10). Because visitors flock to zoos to see
the white tigers, you want to increase the proportion of your
tiger population that is white. How would you go about
doing that?
Your only hope is to alter the population by trying to produce
more animals with the desired white phenotype. In this case,
your best strategy would be to try to breed the white tigers
with each other. Over time, this would lead to the production
of more and more of the white tigers you desire. And as the
generations go by (this will take a while, since the generation
time for tigers is about eight years), your population will
include a higher proportion of white tigers. When this
happens, you will have witnessed evolution, a change in the
allele frequencies of the population.
This example illustrates that evolution doesn’t involve
changing the genetics or physical features of individuals.
Individuals do not evolve. Rather, you change the proportion
of white fur-pigment alleles in the population. An individual
tiger has two alleles for any gene, and within a population, the
proportion of all the alleles coming from any one allele is that
allele’s frequency. It is helpful to think of each allele as having
some “market share” of all of the alleles. In the tiger example,
the white alleles initially had little market share, perhaps as
small as 1%. Over time, though, they came to make up a larger
and larger proportion of the total pigment alleles. And as this
happened, evolution occurred.
Darwin demonstrated that natural selection could be an
efficient mechanism of evolution and a powerful force in
11
Evidence for Evolution
Figure 8-10 Evolution defined. Evolution is a change in allele frequencies in a population.
adapting populations to their environment. Evolution and
natural selection, however, are not the same thing. Natural
selection is one way that evolution can occur, but it is not the
only agent of evolutionary change. It is one of four. They are:
Keeping in mind that evolution is genetic change in a
population, we’ll now explore each of these four forces that
are capable of causing such genetic changes.
1. Mutation
TAKE-HOME MESSAGE
2. Genetic drift
•
Evolution is a change in allele frequencies within a
population. It can occur by four different mechanisms:
mutation, genetic drift, migration, and natural selection.
3. Migration
4. Natural selection
12 8 6
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
8 • 7--------------------------------------------------- Mutation—a direct change in the DNA of an individual—is the
ultimate source of all genetic variation.
of mutations occurring but does not increase the likelihood
In describing the first of the four mechanisms of evolution, it
that a particular mutation is beneficial or detrimental.
is helpful to keep in mind our precise definition of evolution:
a change in the allele frequencies found in a
There are many potential mutation-causing
population. Mutation is an alteration of the
threats in our world today, including tanning
base-pair sequence of an individual’s DNA, and
Tanning beds
beds,
for example, which cause individuals to
when this alteration occurs in a section of the
bombard the
become tan by bombarding the skin with
DNA that codes for a particular gene, the change
body with ultraultraviolet rays similar to those produced by
in the DNA sequence may change the allele. Say,
violet radiation.
the sun. These rays induce tanning just as
for example, that a mutation changes one of the
exposure to the sun does: by stimulating
two blue-eye alleles that an individual possesses
Can they cause
pigment-producing cells to produce more
into a brown-eye allele. If this mutation occurs in
mutations? 8.1
melanin, a pigment that darkens skin color.
the sperm- or egg-producing cells it can be
But because the ultraviolet rays are highpassed on to the next generation; the offspring
energy waves, they can cause mutation just
may be born carrying the brown-eye allele.
as the sun’s rays can. Not surprisingly, researchers have found
When this happens, the proportion of blue-eye alleles in the
that the greater an individual’s exposure to ultraviolet rays,
population, their market share, is slightly reduced and the
the greater the incidence of mutations and, as a consequence,
market share of brown-eye alleles is slightly increased.
of skin cancer.
Evolution has occurred.
Q
Another potential source of mutation-causing energy is of
What causes mutations? For one thing, the incredibly
concern to many more people: mobile phones. The scientific
complex process of cell division can go awry. Duplication of
community is still divided on this question,
the 3 billion base pairs of a cell’s DNA and
however. Although mobile phones do emit
separation of the new and original versions of
Mobile phones
electromagnetic radiation, that radiation is
the DNA into two new cells, after all, are not
sufficiently low level that it is generally
simple processes. Mutations also can be induced
release radiation.
unable to break chemical bonds and therefore
by environmental phenomena. Most
Can
they
cause
cause mutations. The majority of studies have
environmentally induced mutations occur when
brain tumors? 8.2
not found any consistent causal relationship
the DNA of cells is exposed to high-energy
between mobile phone usage and any shortsources or mutation-inducing chemicals (called
or medium-term health hazards.
mutagens). Radioactive isotopes, for example,
emit high-energy particles or photons that can
Since 2004, however, several studies have suggested that
disrupt DNA. This is the reason that radiation therapy is both
further investigation is warranted. A study from 12 laboratories
dangerous and effective as a treatment for cancer: the highfound evidence of DNA damage in cell cultures exposed to
energy source kills most of the cells in its vicinity but can also
levels of radiation similar to those produced by mobile phones.
mutate the DNA in other cells. Ultraviolet radiation, even
Another study found that individuals who had used mobile
from the sun, can cause mutations as well; sunscreen serves as a
phones for 10 years or more had an increased risk of brain
barrier, blocking the harmful radiation from reaching cells and
tumors. A third study discovered an increased risk of brain
the DNA they carry.
tumors and noted that the increased risk was only for the side
of the head on which the phone was used. Stay tuned for
Although environmental factors influence the rate at which
more information on this important question.
mutation occurs, they do not generally influence exactly which
mutations occur.Thus, mutations are random, and whether a
In addition to being an agent of evolutionary change,
mutation happens or not is unrelated to how useful or harmful
mutation has another important role that is relevant to all
that mutation might be.Treatment of some agricultural pest
agents of evolution, including natural selection: mutation is
with harmful chemicals, for example, may increase the number
Q
13
Evidence for Evolution
Figure 8-11 Agents of evolutionary change:
mutation.
DNA may cause the production of a gene product that has
never existed before: instead of blue or brown eyes, the
mutated gene might code for yellow or red. If such a new
allele occurs in the sperm- and egg-producing cells and if it
does not significantly reduce an individual’s “fitness” (which
we’ll consider below), the new allele can be passed on to
offspring and remain in the population. At some future time,
the mutation might even confer higher fitness, in which case
natural selection may cause it to increase in frequency. For this
reason, mutation is critically necessary if natural selection is to
occur: all variation—the raw material for natural selection—
must initially come from mutation.
Despite this vital role in the generation of variation, however,
nearly all mutations reduce an organism’s fitness by causing its
early death or by reducing its reproductive success. Suppose
that you have written a 10-page paper. If you were to
randomly select one letter in the paper and change it to
another letter, is the change more likely to make your paper
better or worse? The answer is obvious: except in very rare
situations (can you imagine one?) the change will hurt your
paper. Similarly, mutations usually change a normally
functioning allele that codes for a normally functioning
protein into an allele that codes for a non-functioning protein.
Almost inevitably, that outcome reduces an organism’s fitness.
For this reason, our bodies are built in ways that protect our
sperm- or egg-producing DNA with a variety of built-in
error-correction mechanisms. As a result, mutations are very,
very rare.
TAKE-HOME MESSAGE
the ultimate source of genetic variation in a population. We
saw that a mutation may lead to the conversion of one allele
to another that is already found within the population, as in
our blue-eye and brown-eye example. More important,
though, a mutation may instead create a completely novel
allele that codes for the production of a new protein (Figure
8-11). That is, a change in the base-pair sequence of a person’s
14 8 7
•
Mutation is an alteration of the base-pair sequence in an
individual’s DNA. Such an alteration constitutes evolution
if it changes an allele that the individual carries. Mutations
can be caused by high-energy sources or chemicals in the
environment and also can appear spontaneously. Mutation
is the only way that new alleles can be created within a
population, and so generates the variation on which natural
selection can act.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
8 • 8--------------------------------------------------- Genetic drift is a random change in allele frequencies
in a population.
Along with mutation, another evolutionary agent is genetic
drift, a random change in allele frequencies. Genetic drift is
best illustrated with an example. Imagine that in a population
there are two alleles for a particular trait, let’s say a cleft chin.
This is a dominant trait, so individuals with either one or two
copies of the dominant allele (CC or Cc) exhibit the cleft
chin (Figure 8-12). Now suppose that two heterozygous (Cc)
people have one child. Which combination of alleles will that child receive? This is impossible to predict because it
depends completely upon which sperm fertilizes which
egg—the luck of the draw. If the couple’s sole child inherited
a recessive allele from each parent, would the population’s
allele frequencies change? Yes. After all, there is now another
individual in the population, and that individual has two
recessive alleles (cc). There are slightly more recessive alleles
in the population. And because a change in allele frequencies
has occurred, evolution has happened. It is equally likely that
this couple’s only child would have received two of the
dominant alleles (CC), rather than the recessive alleles. In
either case, because a change in allele frequencies has
occurred, evolution has happened.
The important factor that distinguishes genetic drift from
natural selection is that the change in allele frequencies is not
related to the alleles’ influence on reproductive success. A cleft
chin does not affect an individual’s ability to reproduce.
This impact of genetic drift is much greater in small
populations than in large populations. In a large population, it
is more likely that any one couple having a child with two
recessive alleles is likely to be offset by another couple having
one child with two dominant alleles. When that occurs, the
Figure 8-12 Agents of evolutionary change: genetic drift. Genetic drift has the greatest impact in
small populations.
15
Evidence for Evolution
Figure 8-13 One way that genetic drift occurs: founder effect.
been evolution. Because the founding members of the new
population will give rise to all subsequent individuals, the new
population will be dominated by the genetic
features that happened to be present in those
Why are
founding fathers and mothers. This type of
Amish people
genetic drift is called the founder effect.
net result is that the overall allele frequencies in the
population have not changed, and evolution has not occurred.
Q
One of the most important consequences of
genetic drift is that it can lead to fixation for
one allele of a gene in a population (see Figure
8-12). Fixation is said to occur when an allele’s
more likely than
frequency in a population reaches 100% (and the
The Amish population in the United States is
other people to
frequency of all other alleles of that gene
believed to have been established by a small
have extra fingers
becomes 0%). If this happens, there is no more
number of founders who happened to carry
and toes?
variability in the population for this gene; all
the allele for polydactyly—the condition of
8.3
individuals will always produce offspring
having extra fingers and toes. As a
carrying only that allele (until new alleles arise
consequence, today this trait, while rare, occurs
through mutation). For this reason, genetic drift reduces the
much more frequently among the Amish than among the rest
genetic variation in a population.
of the U.S. population (Figure 8-13).
Two special cases of genetic drift, the founder effect and
population bottlenecks, are important in the evolution of
populations.
Founder Effect A small number of individuals may leave a
population and become the founding members of a new,
isolated population. The founder population may have
different allele frequencies than the original “source”
population, particularly if the founders are a small sample. If
this new population has different allele frequencies, there has
16 Population Bottlenecks Occasionally, a famine, disease,
or rapid environmental change may cause the deaths of a large
proportion (sometimes as many as 90% or more) of the
individuals in a population. Because the population is quickly
reduced to a small fraction of its original size, this reduction is
called a bottleneck. If the catastrophe is equally likely to
strike any member of the population, the remaining members
are essentially a random, small sample of the original
population. For this reason, the remaining population may not
possess the same allele frequencies as the original population.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
Figure 8-14 Another way that genetic drift occurs: bottleneck effect.
Thus, the consequence of such a population bottleneck would
be evolution through genetic drift (Figure 8-14).
Just such a population bottleneck occurred with cheetahs
near the end of the last ice age, about 10,000 years ago.
Although the cause is unknown—possibly environmental
cataclysm or human hunting pressures—it appears that
nearly all cheetahs died. And although the population has
rebounded, all cheetahs living today can trace their ancestry
back to a dozen or so lucky individuals that survived the
bottleneck. As a result of this past instance of evolution by
genetic drift, there is almost no genetic variation left in the
current population of cheetahs. (And, in fact, a cheetah will
accept a skin graft from any other cheetah, much as identical
twins will from each other.)
TAKE-HOME MESSAGE
8 8
•
Genetic drift is a random change in allele frequencies
within a population, unrelated to the alleles’ influence on
reproductive success. Genetic drift is a significant agent of
evolutionary change primarily in small populations.
17
Evidence for Evolution
8 • 9--------------------------------------------------- Migration into or out of a population may change
allele frequencies.
The third agent of evolutionary change is migration.
Migration, also called gene flow, is the movement of some
individuals of a species from one population to another (Figure 8-15).This movement from population to population
within a species distinguishes migration from the founder
effect, in which individuals migrate to a new habitat, previously
unpopulated by that species. If migrating individuals can
survive and reproduce in the new population, and if they also
carry a different proportion of alleles than the individuals in
their new home, then the recipient population experiences a
change in allele frequencies and, consequently, experiences
evolution. And because alleles are simultaneously lost from the
population that the migrants left, that population, too, will
experience a change in its allele frequencies.
TAKE-HOME MESSAGE
8 9
•
Migration, or gene flow, leads to a change in allele
frequencies in a population as individuals move into or out
of the population.
Figure 8-15 Agents of evolutionary change: migration
(gene flow).
18 CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
8 • 10-------------------------------------------------- When three simple conditions are satisfied, evolution
by natural selection is occurring.
The fourth agent of evolutionary change is natural selection.
This is the mechanism that Darwin identified in The Origin of
Species, in which he noted that three conditions are necessary
for natural selection to occur:
1. There must be variation for the particular trait within a
population.
2. That variation must be inheritable (that is, it must
be capable of being passed on from parents to their
offspring).
3. Individuals with one version of the trait must produce
more offspring than those with a different version of the
trait.
Let’s examine these conditions more closely.
Condition 1: Variation for a Trait Close your eyes and
imagine a dog. What does the dog look like? Chances are, if
50 people were asked this question, we would get descriptions
of 50 different dogs (Figure 8-16): Chihuahuas, Great Danes,
sheepdogs, greyhounds, spaniels, and more. Some are big, some
are small. Some have short hair, some long. They vary in just
about every way that you can imagine. Similarly, if 50 people
were to imagine a human face, a similarly broad range of
images would pop into their heads.Variation is all around us.
Beyond making the world an interesting place to live in,
variation serves another purpose: it is the raw material on
which evolution feeds.
Variation is not limited to physical features such as fur color
and texture or face shape and eye color. Organisms vary in
physiological and biochemical ways, too. Some people can
quickly and efficiently metabolize alcohol, for example.
Others find themselves violently ill soon after sipping a glass
of wine. The same goes for digesting milk. Similarly, we vary
in our susceptibility to poison ivy or diseases such as malaria.
Behavioral variation—from temperament to learning
abilities to interpersonal skills—is dramatic and widespread,
too. So impressed was Darwin with the variation he
observed throughout the world, he devoted the first two
chapters of The Origin of Species to a discussion of
variation in nature and among domesticated animals. Darwin
knew that the variation he saw all around him was an
essential component of evolution by natural selection. He
considered it the first of three conditions necessary for
natural selection.
Condition 2: Heritability The second condition that
Darwin identified as necessary for natural selection was a no
more complex discovery than the first: for natural selection
to act on a trait within a population, offspring must inherit
the trait from their parents. Although inheritance was poorly understood in Darwin’s time, it was not hard to see
that, for many traits, offspring look more like their parents
than like some other random individual in the population
(Figure 8-17). Animal breeders had long known that the
fastest horses generally gave birth to the fastest horses.
Farmers, too, understood that the plants with the highest
productivity generally produced seeds from which highly
productive plants grew. And everyone knew that children
resembled their parents, from their appearance to their
behavior to their temperament. It was enough to know that
Figure 8-16 Necessary conditions for natural selection: 1. Variation for a trait.
19
Evidence for Evolution
deposits. The portly fruit flies have greater reproductive
success than other individuals.
That’s it. Natural selection—certainly one of the most
influential and far-reaching ideas in the history of science—
occurs when three basic conditions are met (Figure 8-19):
1. Variation for a trait
2. Heritability of that trait
3. Differential reproductive success based on that trait
Figure 8-17 Necessary conditions for natural selection: 2. Heritability. Goldie Hawn and daughter Kate Hudson resemble each other.
this similarity between offspring and parents
existed—it was not necessary to understand how
it occurred or to be able to quantify just how
great the similarity was. We call the transmission
of traits from parents to their children through
genetic information inheritance or
heritability.
When these three conditions are satisfied, evolution by natural
selection is occurring. It’s nothing more and nothing less; no
mysterious black box is required. Over time, the traits that lead
some organisms to have greater reproductive success than
others will increase in frequency in a population while traits
that reduce reproductive success will fade away.
Q Most
agricultural pests
evolve resistance
to pesticides.
How does this
happen?
8.4
Condition 3: Differential Reproductive
Success It would be nice to say that Darwin made a
stunning and insightful discovery for the third of the three
conditions necessary for natural selection to occur, but he
didn’t. Rather, he derived the third condition for natural
selection from three fairly simple observations that he made:
Another way of looking at natural selection
is to focus not on the winners (the
individuals who are producing more
offspring), but on the losers. Natural
selection can be turned on its head and
viewed as the elimination of some heritable
traits from a population. If you carry a trait
that makes you a slower-running rabbit, for
example, you are more likely to be eaten by
the fox (Figure 8-20). If running speed is a
heritable trait (and it is), the next generation
1. More organisms are born than can survive.
2. Organisms are continually struggling for existence.
3. Some organisms are more likely to win this struggle
and survive and reproduce. The struggle does not always
involve direct physical contact, but in a world of scarce,
limited resources, finding food or shelter is a zero-sum
game: if one organism is feasting, another is more likely
to be starving.
This three-part observation (which goes by the complex
name differential reproductive success) led Darwin to his
third condition for natural selection: from all the variation
existing in a population, individuals with traits most suited to
reproduction in their environment generally leave more
offspring than do individuals with other traits (Figure 8-18).
For example, in the experiments selecting for fruit flies that
can survive for a long time without food, we saw that the fruit
flies that inherit the ability to pack on the fat when food is
available end up leaving more offspring than those inheriting
a poor ability to pad their little fruit fly frames with fat
20 Figure 8-18 Necessary conditions for natural selection: 3. Differential
reproductive success.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
Figure 8-19 Agents of evolutionary change:
natural selection.
in a population contains fewer slow rabbits. Over time, the
population is changed by natural selection. It evolves.
One of Darwin’s contemporaries, Thomas Huxley, supposedly
cursed himself when he first read The Origin of Species, saying
that he couldn’t believe that he didn’t figure it out on his own.
Each of the three basic conditions is indeed simple and
obvious. The brilliant deduction, though, was to put the three
together and appreciate the consequences.
TAKE-HOME MESSAGE
8 10
•
Natural selection is a mechanism of evolution that occurs
when there is heritable variation for a trait and individuals
with one version of the trait have greater reproductive
success than do individuals with a different version of the
trait. It can also be thought of as the elimination from a
population of alleles that reduce the reproductive rate of
individuals carrying those alleles relative to the reproductive
rate of individuals who do not carry them.
Figure 8-20 Removing the losers. Natural selection can be thought
of as the elimination from a population of traits that confer poor
reproductive success.
21
Evidence for Evolution
4 Through
natural selection,
populations of
organisms can
become adapted to
their environment.
8 • 11--------------------------------------------------- Traits causing some individuals to have more offspring than
others become more prevalent in the population.
“Survival of the fittest.” This is perhaps the most famous
phrase from Darwin’s most famous book. Unfortunately, it can
be a confusing and even misleading phrase. Taken literally, it
seems to be a circular phrase: those organisms that survive
must be the fittest if being fit is defined as the ability to
survive. But we will see that when using the word “fitness” in
discussing evolution, it has little to do with an organism’s
ability to survive or to its physical strength or health. Rather,
fitness has everything to do with an organism’s reproductive
success.
Here’s an interesting side note: the phrase “survival of the
fittest” was coined not by Darwin but by Herbert Spencer, an
influential sociologist and philosopher. Moreover, the phrase
did not appear in Origin of Species when Darwin first
published it. It wasn’t until the fifth edition, 10 years later, that
he used the phrase in describing natural selection.
22 Before we see how fitness affects natural selection and a
population’s adaptation to its environment, let’s define fitness
in a precise way. Fitness is a measure of the relative amount of
reproduction of an individual with a particular phenotype, as
compared with the reproductive output of individuals with
alternative phenotypes. The idea is much easier to understand
with an example. Suppose there are two fruit flies. One fly
carries the genes for a version of a trait that allows it to
survive a long time without food. The other fly has the genes
for a different version of the trait that allows it to survive only
a short while without food. Which fly has the greater fitness?
If the environment is one in which there are long periods of
time without food, such as in the experiment described at the
beginning of the chapter, the fly that can live a long time
without food is likely to produce more offspring than the
other fly and so over the course of its life has greater fitness.
The alleles carried by an individual with high fitness will
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
increase their market share in a population over time and the
population will evolve.
There are three important elements to an organism’s fitness.
1. An individual’s fitness is measured relative to
other genotypes or phenotypes in the population.
Those traits that confer on an individual the highest fitness
will generally increase their market share in a population,
and their increase will always come at the expense of the
market share of alternative traits that confer lower fitness.
2. Fitness depends on the specific environment in
which the organism lives. The fitness value of having
Figure 8-21 An organism’s fitness depends on the environment in
one trait versus another depends on the environment an
which it lives.
organism finds itself in. A sand-colored mouse living in a
beach habitat will be more fit than a chocolatecolored mouse. But that same sand-colored
whether particular traits increase their market
mouse will practically call out to potential
share.
“Survival of
predators when living in the darker brush away
the fittest” is “Survival of the fittest” is a misleading phrase
from the beach. An organism’s fitness, although
since it is the individuals that have the greatest
genetically based, is not fixed in stone and
a misnomer.
reproductive output that are the most fit in any
unchanging—it can change over time and across
Why?
8.5 population. It becomes a more meaningful
habitats (Figure 8-21).
phrase if we consider it as a description of the
3. Fitness depends on an organism’s
fact that the alleles that increase an individual’s
reproductive success compared with other organisms
fitness will “survive” in a population more than the alleles that
in the population. If you carry an allele that gives you
decrease an individual’s fitness.
the trait of surviving for 200 years, but that allele also causes
you to be sterile and incapable of producing offspring,
•
your fitness is zero; that allele will never be passed down to
future generations and its market share will soon be zero.
Fitness is a measure of the relative amount of reproduction
On the other hand, if you inherit an allele that gives you a
of an individual with a particular phenotype, as compared
trait that causes you to die at half the age of everyone else,
with the reproductive output of individuals with alternative
but also causes you to have twice as many offspring as the
phenotypes. An individual’s fitness can vary, depending on
average while you are alive, your fitness is increased. It is
the environment in which the individual lives.
reproductive success that is most important in determining
Q
TAKE-HOME MESSAGE
8 11
23
Evidence for Evolution
8 • 12-------------------------------------------------- Organisms in a population can become better matched
to their environment through natural selection.
If you put a group of humans on the moon, will they flourish?
If you take a shark from the ocean and put it in your swim­
ming pool, will it survive? In both cases, the answer is no.
Organisms are rarely successful when put into novel environ­
ments. And the stranger the new environment, the less likely it
is that the transplanted organism will survive.Why is that?
As Darwin noted over and over again during his travels, the
organisms that possess traits that allow them to better exploit
the environment in which they live will tend to produce
more offspring than the organisms with alternative traits. With
passing generations, a population will be made up of more
and more of these fitter organisms. And, as a consequence,
organisms will tend to be increasingly well matched or
adapted to their environment.
Adaptation refers both to the process by which organisms
become better matched to their environment and to the
specific features that make an organism more fit. Examples of
adaptations abound. Bats have an extremely accurate type of
hearing (called echolocation) for navigating and finding food,
even in complete darkness. Porcupine quills make porcupines
almost impervious to predation (Figure 8-22). Mosquitoes
produce strong chemicals that prevent blood from clotting, so
that they can extract blood from other organisms.
Figure 8-22 Adaptations increase fitness. Quills are an adaptation
that almost completely eliminates the risk of predation for porcupines.
TAKE-HOME MESSAGE
8 12
•
Adaptation—the process by which organisms become better
matched to their environment and the specific features that
make an organism more fit—occurs as a result of natural
selection.
8 • 13-------------------------------------------------- Natural selection does not lead to perfect organisms.
In Lewis Carroll’s Through the Looking-Glass, the Red Queen
tells Alice that “in this place it takes all the running you can
do, to keep in the same place.” She might have been speaking
about the process of evolution by natural selection. After all, if
the least fit individuals are continuously weeded out of the
population, we might logically conclude that, eventually,
fitness will reach a maximum and all organisms in all
populations will be perfectly adapted to their environment.
But this never happens. That is where the Red Queen’s
wisdom comes in.
24 Consider one of the many clearly documented cases of
evolution: that of the beak size of Galápagos finches. Over the
course of a multidecade study, the biologists Rosemary and
Peter Grant closely monitored the average size of the finches’
beaks. They found that the average beak size within a
population fluctuated according to the food supply. During
dry years—when the finches had to eat large, hard seeds—
bigger, stronger beaks became the norm. During wet years,
smaller-beaked birds were more successful since there was a
surplus of small, softer seeds.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
Figure 8-23 What’s fit in one time and place may not be fit in
another. Beak size in Galápagos finches changes along with the
average rainfall.
The ever-changing “average” finch beak illustrates that
2. Variation is needed as the raw material of selection—
adaptation does not simply march toward some optimal
remember, it is the first necessary condition for natural
endpoint (Figure 8-23). Evolution in general, and natural
selection to occur. If a mutation creating a new, “perfect”
version of a gene never occurs, the individuals
selection specifically, do not guide organisms
within a population will never be perfectly
toward “better-ness” or perfection. Natural
adapted. Why, for instance, are there no
selection is simply a process by which, in each
Why doesn’t
mammals
with wheeled appendages? That
generation, the alleles that cause organisms to
natural selection
might be a great trait, but until the genes for
have the traits that make them most fit in that
lead to the proit exist, natural selection cannot increase their
environment tend to increase in frequency.
duction of perfect market share in the population.
If the environment changes, the alleles causing
the traits favored by natural selection may
organisms? 8.6 3. There may be multiple different alleles for
change, too.
a trait, each causing an individual to have the
Q
In the next to last paragraph of Origin of Species,
Darwin wrote: “We may look with some confidence to a
secure future of great length. And as natural selection works
solely by and for the good of each being, all corporeal
[physical] and mental endowments will tend to progress
towards perfection.” In this passage, he overlooks several
different factors that prevent populations from progressing
inevitably toward perfection:
1. Environments change quickly. Natural selection may
be too slow to adapt the organisms in a population to the
constantly moving target that is the environment.
same fitness. In this case, each allele represents
an equally fit “solution” to the environmental challenges.
TAKE-HOME MESSAGE
8 13
•
Natural selection does not lead to organisms perfectly
adapted to their environment because (1) environments
can change more quickly than natural selection can adapt
organisms to them; (2) all possible alleles are not produced
by mutation; (3) there is not always a single optimum
adaptation for a given environment.
25
Evidence for Evolution
8 • 14-------------------------------------------------- Artificial selection is a special case of natural selection.
In practice, plant and animal breeders understood natural
selection before Darwin, they just didn’t know that they
understood it. Farmers bred crops for maximum yield, and dog, horse, and pigeon fanciers selectively bred the
animals with their favorite traits to produce more and more
of the offspring with more and more exaggerated versions of the trait.
The artificial selection used by animal breeders and farmers is
also natural selection because the three conditions are satisfied,
even though the differential reproductive success is being
determined by humans and not by nature. Apple growers, for
example, use artificial selection to produce the wide variety of
apples available: green, yellow, and red, tart and sweet, large
and small. What is important is that it is still differential
reproductive success, and the results are no different. It was a
stroke of genius for Darwin to recognize that the same process
farmers were using to develop new and better crop varieties
was occurring naturally in every population on earth and
always had been.
TAKE-HOME MESSAGE
8 14
•
Animal breeders and farmers are making use of natural
selection when they modify their animals and crops,
because the three conditions for natural selection are
satisfied. Since the differential reproductive success is
determined by humans and not by nature, this type of
natural selection is also called artificial selection.
8 • 15-------------------------------------------------- Natural selection can change the traits in a population
in several ways.
Certain traits are easily categorized—some people have blue
eyes and others have brown eyes, just as some tigers have
white fur and some have orange fur. Other traits, such as
height in humans, are influenced by many genes and
environmental factors so that a continuous range of
phenotypes occurs (Figure 8-24). Whether a given trait is
influenced by one gene or by a complex interaction of many
genes and the environment, it can be subject to natural
selection and be changed in any of several ways.
Directional Selection In directional selection,
individuals with one extreme from the range of variation in
the population have higher fitness. Milk production in cows is
an example. There is a lot of variation in milk production
from cow to cow. As you might expect, farmers select for
breeding those cows with the highest milk production and
have done so for many decades. The result of such selection is
not surprising: between 1920 and 1945, average milk
production increased by about 50% in the United States
(Figure 8-25).
Figure 8-24 Some traits fall into clear categories, others range
continuously.
26 CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
naturally or through artificial selection by breeders.) In
subsequent generations, the average value for the milkproduction trait increases. Note, however, that
the variation for the trait decreases a bit, too,
Turkeys on
because the alleles contributing to one of the
poultry farms
phenotype extremes are eventually removed
from the population. Although the experiment
have such large
has never been done, we can confidently
breast muscles
predict that directional selection for reduced
that they can’t
milk production on these dairy farms would
get close enough have worked equally well.
Q
to each other
to mate. How
can such a trait
evolve?
8.7
Figure 8-25 More milk? Cows have been selected for
their ability to produce more and more milk.
Those at the other end of the range, the cows that produce
the smallest amounts of milk, have reduced fitness, since the
farmers do not allow them to reproduce and therefore
contribute their “less milk” alleles to the subsequent
generation. (This is true whether the selection occurs
Hundreds of experiments in nature and
laboratories demonstrate the power of
directional selection. In fact, it is one of the
marvels of laboratory selection and animal
breeding that nearly any trait they choose to
exaggerate via directional selection responds
dramatically—sometimes with absurd results (Figure 8-26).
Turkeys, for instance, have been selected for increased size of
their breast muscles, which makes them more valuable to
farmers. Directional selection has been so successful that the
birds’ breast muscles are now so large that it has become
impossible for them to mate. All turkeys on large-scale,
industrial poultry farms must now reproduce through artificial
insemination!
Figure 8-26 Patterns of natural selection:
directional selection.
27
Evidence for Evolution
Figure 8-27 Patterns of natural selection: stabilizing selection.
Stabilizing Selection How much did you weigh at birth?
Was it more than 10 pounds? More than 11? Was it close to
the 22½ pounds that Christina Samane’s child weighed when
born in South Africa in 1982? Or was it 5 pounds or less?
Unlike directional selection, for which there is increased
fitness at one extreme and reduced fitness at the other,
stabilizing selection is said to occur when individuals with
intermediate phenotypes are the most fit. The death rate
Figure 8-28 Patterns of natural selection: disruptive selection.
28 CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
among babies, for example, is lowest between
7 and 8 pounds, rising among both lighter and
heavier babies (Figure 8-27). This outcome has
kept the average weight of a baby constant
over many generations, but at the same time the variation in birth weight has decreased as
stabilizing selection has reduced the market
share of those genes associated with high and
low birth weights.
How is medical
Qtechnology
undoing
the work of
natural selection
in optimizing the
number of babies
with normal birth
weights?
8.8
Modern technologies, including Caesarean
deliveries and premature-birth wards, allow
many babies to live who would not have
survived without such technology. This has the unin­tended
consequence of reducing the selection against any alleles
causing those traits, reducing the rate of their removal from
the population.
Disruptive Selection There is a third kind of natural
selection, in which individuals with extreme phenotypes
experience the highest fitness and those with intermediate
phenotypes have the lowest. Although examples of this type
of selection, called disruptive selection, are rare in nature,
the results are not surprising. Among some species of fish—
the Coho salmon, for instance—only the
largest males acquire good territories. They
generally enjoy relatively high reproductive
success. While the intermediate-size fish
regularly get run out of the good territories,
some tiny males are able to sneak in and
fertilize eggs before the territory owner
detects their presence. Consequently, we see
an increase in the frequency of small and large
fish with a reduction in the frequency of
medium-size fish (Figure 8-28).
TAKE-HOME MESSAGE
8 15
•
Acting on multigene traits for which populations show a
large range of phenotypes, natural selection can change
populations in several ways, including directional selection,
in which the average value for the trait increases or
decreases; stabilizing selection, in which the average
value of a trait remains the same while extreme versions
of the trait are selected against; and disruptive selection,
in which individuals with extreme phenotypes have the
highest fitness.
8 • 16-------------------------------------------------- Natural selection can cause the evolution of complex
traits and behaviors.
variation in the rats’ abilities: some rats learned much more
quickly than others how to run the maze. Thompson then
selectively bred the fast learners (called “maze-bright” rats)
with each other and the slow learners (the “maze-dull” rats)
with each other. Over several generations, he developed two
separate populations: rats descended from a line of fast maze
learners and rats descended from a line of slow maze
learners. After only six generations, the maze-dull rats made
twice as many errors as the maze-bright rats
before mastering the maze, while the bright
How can a wing rats were adept at solving complex mazes that
would give many humans some difficulty. Fifty
evolve if 1% of a
years later, it still is unclear which actual genes
wing doesn’t help
are responsible for maze-running behavior, yet
the selection experiment still demonstrates a
an organism fly or
strong genetic component to the behavior.
glide at all?
We have seen that natural selection can change allele
frequencies and modify the frequency with which simple
traits like fur color or turkey-breast size appear in a
population. But what about complex traits, such as behaviors,
that involve numerous physiological and neurological systems?
For instance, can natural selection improve maze-running
ability in rats?
The short answer is yes. Remember, evolution
by natural selection is occurring, changing the
allele frequencies for traits whenever (1) there
is variation for the trait, (2) that variation is
heritable, and (3) there is differential
reproductive success based on that trait. These
conditions can easily be satisfied for complex
traits, including behaviors.
Q
In 1954, to address this question, William
Thompson trained a group of rats to run through a maze for
a food reward (Figure 8-29). There was a huge amount of
8.9
Natural selection can also produce complex traits
in unexpected, roundabout ways. One vexing case involves the
question of how natural selection could ever produce a
29
Evidence for Evolution
Figure 8-29 “Not too complex . . .” Natural selection can produce and alter complex traits,
including behaviors.
Figure 8-30 “Not necessarily a wing . . .” Traits selected for one function may be co-opted by
natural selection for a new function.
30 CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
complex organ such as a fly wing, when 1% or 2% of a wing—
that is, an incomplete structure—doesn’t help an insect to fly. In
other words, while it is clear how natural selection can preserve
and increase traits’ frequencies in populations, how do these
soon-to-be-useful traits increase during the early stages if they
don’t increase the organism fitness at all (Figure 8-30)?
The key to answering this question is that 1% of a wing
doesn’t actually need to function as a wing at all to increase an individual’s fitness. Often, structures appear
because they serve some other purpose. In the case of little
nubs or “almost-wings” on an insect, experiments using
models of insects demonstrated that, as expected, the nubs
confer no benefit at all when it comes to flying. (They don’t
even help flies keep their orientation during a “controlled
fall.”) The incipient wings do help the insects address a
completely different problem, though. They allow much
more efficient temperature control, allowing an insect to
gain heat from the environment when the insect is cold and
to dissipate heat when the insect is hot. Experiments on
heat-control efficiency, in fact, show that as small nubs
become more and more pronounced, they are more and
more effective, probably conferring increased fitness on the
individual in which they occur—but only up to a point.
Eventually, the thermoregulatory benefit stops increasing, even
if the nub length continues to increase. But it is right around
this point that the proto-wing starts to confer some
aerodynamic benefits (see Figure 8-30). Consequently, natural
selection may continue to increase the length of this “almostwing,” but now the fitness increase is due to a wholly different
effect. Such functional shifts explain the evolution of
numerous complex structures that we see today and may be
common in the evolutionary process.
TAKE-HOME MESSAGE
8 16
•
Natural selection can change allele frequencies for genes
involving complex physiological processes and behaviors.
Sometimes a trait that has been selected for one function is
later modified to serve a completely different function.
31
Evidence for Evolution
5 The evidence for the
occurrence of evolution
is overwhelming.
8 • 17-------------------------------------------------- The fossil record documents the process of natural selection.
“It is indeed remarkable that this theory
[evolution] has been progressively accepted by
researchers, following a series of discoveries in
various fields of knowledge. The convergence,
neither sought nor fabricated, of the results of
work that was conducted independently is in itself
a significant argument in favor of this theory.”
— Pope John Paul II, 1996
In the nearly 150 years since Darwin first published The
Origin of Species, thousands of experiments involving his
theory of evolution by natural selection have been
conducted, both in the laboratory and in natural habitats. A wide range of modern methodologies has also been
developed, all contributing to a much deeper understanding
of the process of evolution. This ongoing accumulation of
32 evidence overwhelmingly supports the basic premise that
Darwin put forward, while filling in many of the gaps that
frustrated him.
Here we review the five primary lines of evidence
demonstrating the occurrence of evolution:
1. The fossil record—physical evidence of organisms that
lived in the past
2. Biogeography—patterns in the geographic distribution
of living organisms
3. Comparative anatomy and embryology—growth,
development, and body structures of major groups of
organisms
4. Molecular biology—the examination of life at the
level of individual molecules
5. Laboratory and field experiments—implementation
of the scientific method to observe and study
evolutionary mechanisms
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
Figure 8-31 Evidence for evolution:
the fossil record. Fossils can be used
to reconstruct the appearance of
organisms that lived long ago.
The first of the five lines of evidence is the fossil record.
Although it has been central to much documentation of the
occurrence of evolution, it is a very incomplete record. After
all, the soft parts of an organism almost always decay rapidly
and completely after death. And there are only a few unique
environments (such as tree resin, tar pits, and the bottom of
deep lakes) in which the forces of decomposition are reduced
so that an organism’s hard parts, including bones, teeth, and
shells, can be preserved for thousands or even millions of years.
These remains, called fossils, can be used to reconstruct what
or­ganisms must have looked like long ago. Such
reconstructions often provide a clear record of evolutionary
change (Figure 8-31).
The use of radiometric dating helps in further painting a
picture of organisms’ evolutionary history by telling us the age
of the rock in which a fossil has been found (Figure 8-32). In
Darwin’s time, it was assumed that the deeper down in the
earth a fossil was found, the older it was. Radiometric dating
goes a step further, making it possible to determine not just
the relative age of fossils, but also their absolute age. This is
accomplished by evaluating the amounts of certain radioactive
isotopes present in fossils. Radioactive isotopes in a rock begin
breaking down into more stable compounds as soon as the
rock is formed, and they do so at a constant rate. Nothing can
alter this. By measuring the relative amounts of the radioactive
isotope and the leftover decay product in the rock where a
fossil is found, the age of the rock and thus of the fossil can be
calculated.
Figure 8-32 How old is that fossil? Radiometric dating of uranium 238
helps to determine the age of rocks and the fossils in them.
33
Evidence for Evolution
Radiometric dating confirms that the earth is very old. Rocks more than 3.5 billion years old have been found on all
of the earth’s continents, with the oldest so far found in
north­western Canada. By using the radioactive isotope
uranium 238, with a half-life of 4.5 billion years, researchers
have determined that the earth is about 4.6 billion years old
and that the earliest organisms appeared at least 3.8 billion
years ago. Radiometric dating also makes it possible to put the
fossil record in order. By dating all the fossils discovered in one
locale, paleontologists can learn if and how the organisms
were related to each other and how groups of organisms
changed over time.
Paleontologists must deal with the fact, however, that fossil
formation is an exceedingly unlikely event, and when it does
occur it represents only those organisms that happened to
live in that particular area and that also had physical
structures that could leave fossils. For this reason, the fossil
record is annoy­ingly incomplete. Entire groups of organisms
have left no fossil record at all and some others have
numerous gaps. Still, fossils have been found linking all of
the major groups of vertebrates.
The evolutionary history of horses is among the most well
preserved in the fossil record. First appearing in North
America about 55 million years ago, horses then radiated
around the world, with more recent fossils appearing in
Eurasia and Africa. These fossils exhibit distinct adaptations
to those environments. Later, about 1.5 million years ago,
much of the horse diversity—including all North American
horse species—disappeared, leaving only a single remaining
genus, or group of species, called Equus. Because there is
now only one horse genus on earth, it is tempting to
imagine a simple linear path from modern horses straight
back through their 55-million-year evolutionary history. But
that’s just not how evolution works. In reality, there have
been numerous branches of horses that have split off over
evolutionary time, flourished for millions of years, and only
recently gone extinct. What we see living today is only a
single branch of a greatly branched evolutionary tree (Figure 8-33).
The fossil record provides another valuable piece of evidence
for the process of natural selection in the form of “missing
links.” These are fossils that demonstrate a link between groups
of species believed to have shared a common ancestor. One
such “missing link” is Tiktaalik (Figure 8-34). First found in
northern Canada and estimated to be 375 million years old,
Tiktaalik fossils seem to represent a transitional phase between
fish and land animals.These creatures, like fish, had gills, scales,
Figure 8-33 An evolutionary family tree. The branching evolutionary tree of the horse.
34 CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
Figure 8-34 A missing link? Tiktaalik
seems to be a transitional phase between
fish and land animals.
and fins, but they also had arm-like joints in their fins and
could drag their bodies across land in much the same way as a
marine mammal such as a seal.
TAKE-HOME MESSAGE
8 17
•
Radiometric dating confirms that the earth is very old and
makes it possible to determine the age of fossils. Analysis
of fossil remains enables biologists to reconstruct what
organisms looked like long ago, learn how organisms
were related to each other, and understand how groups of
organisms evolved over time.
8 • 18-------------------------------------------------- Geographic patterns of species distributions reflect their
evolutionary histories.
The study of the distribution patterns of living organisms
around the world is called biogeography. This is the second
line of evidence that helps us to see that evolution occurs and
to better understand the process.The patterns of biogeography
that Darwin and many subsequent researchers have noticed
provide strong evidence that evolutionary forces are responsible
for these patterns. Species often more closely resemble other
species that live less than a hundred miles away but in radically
different habitats than they do species that live thousands of miles
away in nearly identical habitats. On Hawaii, for example, nearly
every bird is some sort of modified honeycreeper (a small, finchlike bird).There are seed-eating honeycreepers, woodpecker
honeycreepers, and curved-bill nectar-feeding honeycreepers.
There are even parrot honeycreepers (Figure 8-35).
Figure 8-35 Evidence for evolution: biogeography. The Hawaiian honeycreepers resemble a common
ancestor from mainland North America, but all have unique features.
35
Evidence for Evolution
When it comes to species distributions, history matters. Species
were not designed from scratch to fill a particular niche.
Rather, whatever arrived first—usually a nearby species—took
up numerous different lifestyles in numerous different habitats,
and the populations ultimately adapted to and evolved in each
environment. In Hawaii, it seems that a finch-like descendant
of the honeycreepers arrived 4–5 million years ago and rapidly
evolved into a large number of diverse species.The same
process has occurred and continues to occur in all locales, not
just on islands.
Large isolated habitats also have interesting biogeographic
patterns. Australia and Madagascar are filled with unique
organisms that are clearly not closely related to organisms
elsewhere. In Australia, for example, marsupial species, rather
than placental mammals, fill all of the usual roles. There are
marsupial “wolves,” marsupial “mice,” marsupial “squirrels,”
and marsupial “anteaters” (Figure 8-36). They physically
resemble their placental counterparts for most traits, but
molecular analysis shows that they are actually more closely
related to each other, sharing a common marsupial ancestor.
Their relatedness to each other is also revealed by similarities
in their reproduction: females give birth to offspring at a
relatively early state of development and the offspring finish
their development in a pouch. The presence of marsupials in
Australia does not simply mean that marsupials are better
adapted than placentals to the Australian habitat. When
placental organisms have been transplanted to Australia they
do just fine, often thriving to the point of endangering the
native species.
A good designer would use the best designs over and over
again wherever they might fit. Biogeographic patterns such as
those seen in honeycreepers and in the marsupials of Australia
illustrate that evolution, unlike a good designer, is more of a
tinkerer. Evolution takes whatever populations are at hand in a
particular location then gradually changes their traits, and the
species become better adapted to the habitats they occupy.
TAKE-HOME MESSAGE
8 18
•
Observing geographic patterns of species distributions,
particularly noting similarities and differences among
species living close together but in very different habitats
and among species living in similar habitats but located far
from one another, helps us to understand the evolutionary
histories of populations.
Figure 8-36 Evidence for evolution: biogeography. Many Australian marsupials resemble
placental counterparts, though they are not closely related.
36 CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
8 • 19-------------------------------------------------- Comparative anatomy and embryology reveal common
evolutionary origins.
If you observe any vertebrate embryo, you will see that it
passes through a stage in which it has little gill pouches on the
sides of the neck. It will also pass through a stage in which it
possesses a long bony tail. This is true whether it is a human
embryo or that of a turtle or a chicken or a shark. These gill
pouches disappear before birth in all but the fishes. Similarly,
we don’t actually find humans with tails. Why do they exist
during an embryo’s development? Such common
embryological stages indicate that the organisms share a
common ancestor, from which all have been modified (Figure 8-37). Study of these developmental stages and the
adult body forms of organisms provides our third line of
evidence, helping us to see that evolution occurs and to better
understand the process.
Among adults, several features of anatomy reveal the ghost of
evolution in action. We find, for example, that many related
organisms show unusual similarities that can be explained only
through evolutionary relatedness. The forelimbs of mammals
are used for a variety of functions in bats, porpoises, horses,
and humans (Figure 8-38). If each had been designed
specifically for the uses necessary to that species, we would
expect dramatically different designs. And yet in each of these
species, we see the same bones—modified extensively—that
betray the fact that they share a common ancestor. These
features are called homologous structures.
Figure 8-37 Evidence for evolution: embryology. Structures derived
from common ancestry can be seen in embryos.
Figure 8-38 Evidence for evolution: comparative anatomy. Homologous bone structures among some mammals.
37
Evidence for Evolution
Q
At the extreme, homologous
structures sometimes come
The human
to have little or no function
appendix serves
at all. Such evolutionary
no function. Why
leftovers, called vestigial
structures, exist because
are we all born
they had value ancestrally.
with one?
8.10
Some vestigial structures in
mammals include the molars that continue to
grow in vampire bats, even though these bats consume a
completely liquid diet (Figure 8-39); eye sockets (with no
eyes) in some populations of cave-dwelling fish; and pelvic
bones in whales that are attached to nothing (but serve as an
important attachment point for leg bones in nearly all other
mammals). Even humans have a vestigial organ—the
appendix. It is greatly enlarged in our relatives the great apes,
in whom it hosts cellulose-digesting bacteria that aid in
breaking down the plants in their diet, but in humans it
serves no purpose.
Not all organisms with adaptations that appear similar actually
share ancestors. We see flying mammals (bats) and flying
insects (locusts) (Figure 8-40). Similarly, dolphins and penguins
live in similar habitats and have flippers that help them to
swim. In both examples, however, all of the analogous
Figure 8-40 Evidence for evolution: convergent evolution and
analogous structures.
structures developed from different original structures. Natural
selection—in a process called convergent evolution—uses
the different starting materials available (such as a flipper or a
forelimb) and modifies them until they serve similar purposes,
much as we saw in the marsupial and placental mammals in
Figure 8-36.
TAKE-HOME MESSAGE
8 19
•
Similarities in the anatomy of different groups of organisms
and in their physical appearance as they proceed through
their development can reveal common evolutionary origins.
Figure 8-39 Evidence for evolution: vestigial structures.
38 CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
8 • 20-------------------------------------------------- Molecular biology reveals that common genetic
sequences link all life forms.
The development of new technologies for deciphering and
comparing the genetic code provides our fourth line of
evidence that evolution occurs. First and foremost is the fact
that all living organisms share the same genetic code. From
microscopic bacteria to flowering plants to insects and
primates, the molecular instructions for building organisms
and transmitting hereditary information are the same: four
simple bases, arranged in an almost unlimited variety of
sequences. This discovery strongly suggests that all living
organisms are related.
When we examine the similarity of DNA among related
individuals within a species, we find that they share a greater
proportion of their DNA than do unrelated individuals. This
is not unexpected; you and your siblings got all of your DNA
from the same two parents, while you and your cousins each
got half of your DNA from the same two grandparents. The
more distantly you and another individual are related, the
more your DNA differs.
We can measure the DNA similarity between two species by
comparing their DNA sequences for individual genes. For
example, let’s look at the gene that codes for the amino acids
used to build the hemoglobin molecule. In vertebrate animals,
hemoglobin is found inside red blood cells, where its function
is to carry oxygen throughout the body. It is made of two
chains of amino acids, the alpha chain and the beta chain. In
humans, the beta chain has 146 amino acids. In rhesus mon­keys,
this beta chain is nearly identical: of the 146 amino acids, 138
are the same as those found in human hemoglobin, and only 8
are different. In dogs the sequence is still similar, but a bit less so,
with 32 different amino acids.When we look at non-mammals
such as birds, we see about 45 amino acid differences, and
between humans and lamprey eels (still verte­brates, but it has
been more than 450 million years since we shared a common
ancestor), there are 125 amino acid differences.
The differences in the amino acid structure of the beta
hemoglobin chain (and remember that this structure is
governed by an allele or alleles of a particular gene) seem to
indicate that humans have more recently shared a common
ancestor with rhesus monkeys than with dogs. And that we
have more recently shared an ancestor with dogs than with
birds or lampreys. These findings are just as we would expect,
based on estimates of evolutionary relatedness made from
comparative anatomy and embryology as well as those based
on the fossil record. It is as if there is a molecular clock that is
Figure 8-41 An evolutionary clock? Genetic similarities (and
differences) demonstrate species relatedness.
ticking. The longer two species have been evolving on their
own, the greater the number of changes in amino acid
sequences—or “ticks of the clock” (Figure 8-41).
TAKE-HOME MESSAGE
8 20
•
All living organisms share the same genetic code. The
degree of similarity in the DNA of different species can
reveal how closely related they are and the amount of time
that has passed since they last shared a common ancestor.
39
Evidence for Evolution
8 • 21-------------------------------------------------- Laboratory and field experiments enable us to watch
evolution in progress.
A fifth line of evidence for the occurrence of evolution comes
from multigeneration experiments and observations. Until
recently, people thought that evolution was too slow a process
to be observed in action. By choosing the right species—
preferably organisms with very short life spans—and designing
careful experiments, however, it turns out that it is actually
possible to observe and measure evolution as it is happening.
infections are resistant to these antibiotics. The meaning of
such unintentional natural selection “experiments” is clear and
consistent: evolution is occurring all around us.
And finally, let’s return to the spectacularly starvation-resistant
flies introduced at the beginning of this chapter. In them we
saw an unambiguous demonstration that natural selection can
occur, that it has the potential to produce dramatic changes in
In one clever study, researchers studied populations of grass on
golf courses—a habitat where lawnmower blades represent a
significant source of mortality.They realized that although all of
the grass on golf courses was the same species, on the putting
greens it was cut very frequently, whereas on the fairways it was
cut only occasionally, and in the rough it was almost never cut
at all (Figure 8-42). Over the course of only a few years,
significant changes occurred in these different grass populations.
The grass plants on the greens came to be short lived, with
rapid development to reproductive age and very high seed
output. For the plants in these populations, life was short and
reproduction came quickly. (For those in which it did not
come quickly, it did not come at all; hence their lack of
representation in the population.) Plants on the fairways had
slightly slower development and reduced seed output, while
those in the rough had the slowest development and the lowest
seed output of all.When plants from each of the habitats were
collected and grown in greenhouses, the dramatic differences in
growth, development, and life span remained, confirming that
there had been changes in the frequency of the various alleles
controlling the traits of life span and reproductive output in the
three populations; in other words, evolution had occurred.
A more disturbing line of evidence for the occurrence of
evolution in nature comes from the evolution of antibioticresistant strains of bacteria that cause illness in humans. For
example, in the 1940s when penicillin was first used as a
treatment for bacterial infections, it was uniformly effective in
killing Staphylococcus aureus. Today, more than 90% of isolated
S. aureus strains are resistant to penicillin (Figure 8-43). Because
penicillin has become such a pervasive toxin in the
environment of Staphylococcus, natural selection has led to an
increase in the frequency of the alleles that make these strains
resistant. As a consequence, humans are increasingly at risk for
becoming infected with Staphylococcus and getting diseases
such as pneumonia and meningitis. In the 1960s, the
antibiotics methicillin and oxacillin were developed. Like
penicillin, they initially had nearly complete effectiveness
against Staphylococcus. Today, though, nearly a third of staph
40 Figure 8-42 Evolution in progress: grasses. Different mowing
patterns can cause evolution in the grass on a golf course.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
physical features and physiological processes, and that it can
bring about these changes very quickly. And, perhaps most
important, we saw that replicating the same evolutionary
process over and over repeatedly produced the same
predictable results. In the laboratory, the farm, the doctor’s
office, the bathroom sink, the deserts, streams, and forests,
evolution is occurring.
Reflecting on both the process and products of evolution and
natural selection, in the final paragraph of The Origin of Species
Darwin eloquently wrote: “There is grandeur in this view of
life . . . and that . . . from so simple a beginning endless forms
most beautiful and most wonderful have been, and are being,
evolved.”
TAKE-HOME MESSAGE
Figure 8-43 Evolution in progress: disease-causing bacteria.
Antibiotic resistance has evolved in Staphylococcus.
8 21
•
Multiply replicated, controlled laboratory selection
experiments and long-term field studies of natural
populations—including observations on antibiotic-resistant
strains of disease-causing bacteria—enable us to watch and
measure evolution as it occurs.
41
Evidence for Evolution
streetBio
Knowledge You Can Use
Did you know? Antibacterial soaps may be
dangerous. And it may be your fault. Fact: The
antimicrobial agent, called triclosan, in most
antibacterial soaps takes several minutes to kill
bacteria.
Q: How long do you wash your hands? On average, people
spend less than a minute washing their hands.
Q: If you’re not killing all of the bacteria, which ones
are you killing? Some bacteria are easily killed by the
antimicrobial chemical in soaps. Others take longer. In the
first minute you’re killing only the weak bacteria (and not all of
the 800,000 to 1,000,000 bacteria residing on your hands).
Q:
After one minute, what is left? After a minute of handwashing, only the strong bacteria—those that happen
to carry triclosan-resistant genes—remain and continue
reproducing. (And the next time you wash your hands, you may
pick up those super-strength drug-resistant bacteria from the sink
area.) You’re causing evolution to happen. (And you may not
want to.)
Q: What do you make of this? What can you do? The two
simplest alternatives are to wash your hands for much longer or to
stop using soaps with antibacterial agents in them.
42 CHAPTER 8 • EVOLUTION AND NATURAL SELECTION

Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
The characteristics of individuals in a
population can change over time. We can
observe such change in nature and can
even cause such change to occur.
2 Darwin journeyed to a new idea.
Following years of observations on
worldwide patterns of the living
organisms and fossils of plant and animal
species, Charles Darwin developed a
theory of evolution by natural selection
that explained how populations of species can change
over time.
3 Four mechanisms can give rise to evolution.
Evolution—a change in the allele
frequencies in a population—can occur
via genetic drift, migration, mutation, or
natural selection.
4 Through natural selection, populations
reproductive output. The alleles that yield traits that
confer the highest fitness on the individuals carrying
them will increase their market share in a population
over time, but this process does not necessarily lead
to “perfect” organisms in an environment. Artificial
selection, in which humans determine which individuals
will have the highest reproductive success, is a type of
natural selection. Directional, stabilizing, and disruptive
selection can alter the average value of a trait and/
or the variation for a trait in a population. Natural
selection also can produce and modify complex traits,
including behaviors. This modification may occur as
traits selected for one purpose later confer increased
fitness for a different purpose.
5 The evidence for the occurrence of
evolution is overwhelming.
Many overwhelming lines of evidence
document the occurrence of evolution
and point to the central and unifying
role of evolution by natural selection
in helping us to better understand all
other ideas and facts in biology.
of organisms can become adapted to their
environment.
When there is variation for a trait, and
the variation is heritable, and there is
differential reproductive success based on
that trait, evolution by natural selection is
occurring. Fitness is an individual’s relative
big ideas in evolution and natural selection
1 Evolution is an ongoing process.
key terms---------------------------------------------------------------------------------------------------------adaptation, p. 24
biogeography, p. 35
bottleneck effect, p. 16
convergent evolution, p. 38
differential reproductive success, p. 20
directional selection, p. 26
disruptive selection, p. 29
evolution, p. 3
fitness, p. 23
fixation, p. 16
fossil, p. 33
founder effect, p. 16
gene flow, p. 18
genetic drift, p. 15
heritability, p. 20
homologous structure, p. 37
inheritance, p. 20
migration, p. 18
mutagen, p. 13
mutation, p. 13
natural selection, p. 3
population, p. 2
radiometric dating, p. 33
stabilizing selection, p. 28
trait, p. 8
vestigial structure, p. 38
43
Evidence for Evolution
Check Your Knowledge---------------------------------------------------------- 1. The average time to death from starvation in a fruit fly is
about 20 hours. Selecting for increased starvation resistance in
fruit flies
a) has no effect because starvation resistance is a not a trait
that influences fruit fly fitness.
b) has little effect because ongoing mutation continuously
reduces starvation resistance, counteracting any benefits
from selection.
c) cannot increase their survival time because there is no
genetic variation for this trait.
d) has no effect because starvation resistance is too complex a
trait, dependent on the effects of too many genes.
e) can produce populations in which the average time to
death from starvation is 160 hours.
2. Georges Cuvier’s discovery of fossils of Irish elk and giant
ground sloths
a) demonstrated that extinction must occur.
b) was possible because of Buffon’s determination that the
earth was more than 6,000 years old.
c) was possible only following Darwin’s publication of The
Origin of Species.
d) was made in deep ocean trenches.
e) suggested that species were immutable.
3. While on the voyage of the HMS Beagle, Darwin
a) wrote The Origin of Species.
b) nurtured his love of studying nature, exploring plant and
animal diversity, and collecting fossils.
c) discovered a love for sea travel.
d) studied Malthus’s book Essay on the Principle of Population.
e) corresponded extensively with Alfred Russel Wallace about
their ideas on evolution.
4. Which one of the following statements BEST describes the
difference between artificial and natural selection?
a) Natural selection is limited to physical traits and artificial
selection is not.
b) Artificial selection has produced many of the most
delicious food items for humans; natural selection has not.
c) Natural selection acts without the input of humans;
artificial selection requires human input.
d) Charles Darwin understood natural selection but was
unaware of artificial selection in his time.
e) Natural selection works on all species; artificial selection
works only on laboratory-raised species.
5. Which of the following statements about Charles Darwin is
FALSE?
a) He spent five years traveling the world observing living
organisms and collecting fossils.
b) He was under constant pressure from his father to make
something of himself.
c) He dropped out of medical school.
d) He was enthusiastic about unleashing his theory of natural
selection on the world as soon as he thought of it.
e) He and Alfred Russel Wallace independently came up with
the theory of evolution by natural selection.
44 6. Evolution occurs
a) only when the environment is changing.
b) only through natural selection.
c) almost entirely because of directional selection.
d) only via natural selection, genetic drift, migration, or
mutation.
e) by altering physical traits but not behavioral traits.
7. Which of the following statements about mutations is NOT
true?
a) Mutations are almost always random with respect to the
needs of the organism.
b) A mutation is any change in an organism’s DNA.
c) Most mutations are harmful or neutral to the organism in
which they occur.
d) The origin of genetic variation is mutation.
e) All of these statements are true.
8. The chief concern among conservation biologists trying to
protect small populations is
a) to preserve genetic diversity.
b) that mutation rates are much higher in small populations.
c) that natural selection can only operate in large populations.
d) to maximize rates of allele fixation.
e) to reduce the duration of genetic bottlenecks.
9. When a group of individuals colonizes a new habitat, the
event is likely to be an evolutionary event because
a) members of a small population have reduced rates of
mating.
b) gene flow increases.
c) mutations are more common in novel environments.
d) new environments tend to be inhospitable, reducing
survival there.
e) small founding populations are rarely genetically
representative of the initial population.
10. To establish that evolution by natural selection is operating
in a population, one must demonstrate variability for a trait,
heritability of that trait, differential reproductive success based
on that trait, and
a) increased complexity of the organism.
b) random mating.
c) progress.
d) continuous change in the environment.
e) nothing else.
11. “Survival of the fittest” may be a misleading phrase to
describe the process of evolution by natural selection because
a) it is impossible to determine the fittest individuals in
nature.
b) survival matters less to natural selection than reproductive
success does.
c) natural variation in a population is generally too great to
be influenced by differential survival.
d) fitness has little to do with natural selection.
e) reproductive success on its own does not necessarily
guarantee evolution.
CHAPTER 8 • EVOLUTION AND NATURAL SELECTION
Evolution Is Ongoing
Darwin’s Journey
Four Mechanisms
Natural Selection
12. Adaptation
a) refers both to the process by which organisms become
better matched to their environment and to the features of
an organism that make it more fit than other individuals.
b) cannot occur in environments influenced by humans.
c) is possible only when there is no mutation.
d) is responsible for the fact that porcupines are at unusually
high risk of predation.
e) occurs for physical traits but not behaviors.
13. Adaptations shaped by natural selection
a) are magnified and enhanced through genetic drift.
b) are unlikely to be present in humans living in industrial
societies.
c) may be out of date, having been shaped in the past under
conditions that differed from those in the present.
d) represent perfect solutions to the problems posed by
nature.
e) are continuously modified so that they are always fitted to
the environment in which an organism lives.
14. Artificial selection was used on corn to produce a single strain
of corn with increased growth rates and greater resistance to
a fungus. Although farmers have continued to select for these
traits, the productivity of this strain is no longer increasing.
This suggests that
a) the population size has been decreasing.
b) all or most of the natural variation for these traits has been
eliminated.
c) gene migration is a major evolutionary agent in corn.
d) long-term disruptive selection may lead to speciation.
e) artificial selection is not as strong as natural selection.
15. In a population in which a trait is exposed to stabilizing
selection over time
a) neither the average nor the variation for the trait changes.
b) both the average value and variation for the trait increase.
c) the average value increases or decreases and the variation
for the trait decreases.
d) the average value for the trait stays approximately the same
and the variation for the trait decreases.
e) the average value for the trait stays approximately the same
and the variation for the trait increases.
16. Maze-running behavior in rats
a) is too complex a trait to be influenced by natural selection.
b) is a heritable trait.
c) is not influenced by natural selection because it does not
occur in rats’ natural environment.
d) shows no variation.
e) is influenced primarily by mutation in the laboratory.
17. A fossil is defined most broadly as
a) the preserved pieces of hard parts (e.g., shell or bone) of
extinct animals.
b) any preserved remnant or trace of an organism from the
past.
c) the preserved bones of vertebrates.
d) a piece of an organism that has turned into rock.
e) the process of preservation of intact animal bodies.
18. Which of the following are true about marsupial mammals?
a) fill many niches in Australia that are occupied by placental
mammals in other parts of the world.
b) are less fit than placental mammals.
c) have gone extinct as a result of the greater fitness of
placental mammals.
d) are more closely related to each other than they are to
placental mammals.
e) Both a) and d) are correct.
19. Convergent evolution can occur only when two species
a) have a recent common ancestor.
b) live in the same geographic area.
c) are separated by a barrier such as a new river.
d) evolve under similar selective forces.
e) are both unpalatable to predators.
20. Which of the following can be used to calibrate a “molecular
clock” for a phylogenetic tree constructed from DNA
sequences?
a) the fossil record
b) geological events
c) the mutation rate of other genes
d) Both a) and b) can be used for calibration.
e) All of the above can be used for calibration.
21. Evolution
a) occurs too slowly to be observed in nature.
b) can occur in the wild but not in the laboratory.
c) is responsible for the increased occurrence of antibioticresistant bacteria.
d) does not occur in human-occupied habitats.
e) None of these statements is correct.
Short-Answer Questions
1. Distinguish between evolution and natural selection. (Restrict
your answer to 30 words or fewer.)
2. What is genetic drift? Why is it a more potent agent of
evolution in small populations rather than large populations?
3. In The Origin of Species, Charles Darwin wrote: “We may look
with some confidence to a secure future of great length. And
as natural selection works solely by and for the good of each
being, all corporeal and mental endowments will tend to
progress towards perfection.” Describe three reasons why he
is wrong.
4. How does the increasing frequency of antibiotic-resistant
strains of bacteria represent an example of the occurrence
evolution?
See Appendix for answers. For additional study questions,
go to www.prep-u.com.
45
Evidence for Evolution

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