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12/7/2016
CAMPBELL
BIOLOGY
Endless Forms Most Beautiful
TENTH
EDITION
Reece • Urry • Cain • Wasserman •
Minorsky • Jackson
21
How
Evolution
Works
 A new era of biology began in 1859 when Charles
Darwin published The Origin of Species
 The Origin of Species focused biologists’ attention
on the great diversity of organisms
Lecture Presentation by
Nicole Tunbridge and
Kathleen Fitzpatrick
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.1
 Darwin noted that current species are
descendants of ancestral species
 Evolution can be defined by Darwin’s phrase
descent with modification
 Evolution can be viewed as both a pattern and a
process
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.1a
Concept 21.1: The Darwinian revolution
challenged traditional views of a young Earth
inhabited by unchanging species
 Darwin’s ideas had deep historical roots
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.2
Figure 21.2a
1809
Lamarck publishes his
hypothesis of evolution.
1809
Lamarck publishes his
hypothesis of evolution.
1798
Malthus publishes
“Essay on the Principle
of Population.”
1795
Hutton proposes
his principle of
gradualism.
1790
1809
Charles Darwin
is born.
Sketch of a flying
frog by Wallace
1812
Cuvier publishes his
extensive studies of
vertebrate fossils.
1830
Lyell publishes
Principles of Geology.
1831–1836
Darwin travels
around the world
on HMS Beagle.
1858
While studying species in the
Malay Archipelago, Wallace
(shown above in 1848) sends
Darwin his hypothesis of
natural selection.
1870
1859
On the Origin of
Species is published.
1844
Darwin writes his
essay on descent
with modification.
1798
Malthus publishes
“Essay on the Principle
of Population.”
1795
Hutton proposes
his principle of
gradualism.
Marine iguana
in the
Galápagos
Islands
© 2014 Pearson Education, Inc.
1790
1809
Charles Darwin is born.
© 2014 Pearson Education, Inc.
1812
Cuvier publishes his
extensive studies of
vertebrate fossils.
12/7/2016
Figure 21.2b
Figure 21.2c
1830
Lyell publishes
Principles of Geology.
1831–1836
Darwin travels
around the world
on HMS Beagle.
1858
While studying species in the
Malay Archipelago, Wallace
sends Darwin his hypothesis
of natural selection.
1870
1859
On the Origin of
Species is published.
1812
Cuvier publishes his extensive
studies of vertebrate fossils.
1844
Darwin writes his
essay on descent
with modification.
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.2d
Figure 21.2e
Marine iguana
in the
Galápagos
Islands
Sketch of a flying
frog by Wallace
1831–1836
Darwin travels around the world on
HMS Beagle.
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.2f
Figure 21.2g
1858
While studying species
in the Malay Archipelago,
Wallace (shown above in
1848) sends Darwin his
hypothesis of natural
selection.
© 2014 Pearson Education, Inc.
1859
On the Origin of Species
is published.
© 2014 Pearson Education, Inc.
Scala Naturae and Classification of Species
 The Greek philosopher Aristotle viewed species as
fixed and arranged them on a scala naturae
 The Old Testament holds that species were
individually designed by God and therefore perfect
 Carolus Linnaeus interpreted organismal
adaptations as evidence that the Creator had
designed each species for a specific purpose
 Linnaeus was the founder of taxonomy, the branch
of biology concerned with classifying organisms
 He developed the binomial format for naming
species (for example, Homo sapiens)
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.3
Ideas About Change over Time
 The study of fossils helped to lay the groundwork
for Darwin’s ideas
Sedimentary rock
layers (strata)
 Fossils are remains or traces of organisms from
the past, usually found in sedimentary rock, which
appears in layers called strata
Younger stratum
with more recent
fossils
Older stratum
with older fossils
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Video: Grand Canyon
 Paleontology, the study of fossils, was largely
developed by French scientist Georges Cuvier
 Cuvier speculated that the boundaries between
strata represent catastrophic events
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Lamarck’s Hypothesis of Evolution
 Geologists James Hutton and Charles Lyell
perceived that changes in Earth’s surface can
result from slow continuous actions still operating
today, and at the same rate
 This view strongly influenced Darwin’s thinking
© 2014 Pearson Education, Inc.
 Lamarck hypothesized that species evolve through
use and disuse of body parts and the inheritance
of acquired characteristics
 The mechanisms he proposed are unsupported by
evidence
© 2014 Pearson Education, Inc.
Figure 21.4
Concept 21.2: Descent with modification by
natural selection explains the adaptations of
organisms and the unity and diversity of life
 Some doubt about the permanence of species
preceded Darwin’s ideas
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Darwin’s Research
The Voyage of the Beagle
 As a boy and into adulthood, Charles Darwin had
a consuming interest in nature
 During his travels on the Beagle, Darwin collected
specimens of South American plants and animals
 Darwin first studied medicine (unsuccessfully), and
then theology at Cambridge University
 He observed that fossils resembled living species
from the same region, and living species
resembled other species from nearby regions
 After graduating, he took an unpaid position as
naturalist and companion to Captain Robert
FitzRoy for a 5-year around the world voyage on
the Beagle
© 2014 Pearson Education, Inc.
 He experienced an earthquake in Chile and
observed the uplift of rocks
© 2014 Pearson Education, Inc.
Figure 21.5
 Darwin was influenced by Lyell’s Principles of
Geology and thought that the earth was more than
6,000 years old
 His interest in geographic distribution of species
was kindled by a stop at the Galápagos Islands
west of South America
 He hypothesized that species from South America
had colonized the Galápagos and speciated on
the islands
© 2014 Pearson Education, Inc.
Darwin in 1840,
after his return
from the
voyage
HMS Beagle in port
Great
Britain
NORTH
AMERICA
The
Galápagos
Islands
ATLANTIC
OCEAN
Pinta
Santiago
Isabela
0
20 40
AFRICA
PACIFIC
OCEAN
SOUTH
AMERICA
Genovesa
Marchena
Fernandina
EUROPE
Equator
Daphne
Islands
Pinzón
Santa Santa
Cruz
Fe
Florenza
Kilometers
© 2014 Pearson Education, Inc.
San
Cristobal
Española
Chile
PACIFIC
OCEAN
Equator
Brazil
Cape of
Good Hope
Argentina
Cape Horn
Malay
Archipelago
PACIFIC
OCEAN
AUSTRALIA
Tasmania
New
Zealand
12/7/2016
Figure 21.5a
Figure 21.5b
Great
Britain
NORTH
AMERICA
EUROPE
ATLANTIC
OCEAN
AFRICA
SOUTH
AMERICA
Equator
Malay
Archipelago
Brazil
Chile
PACIFIC
OCEAN
PACIFIC
OCEAN
AUSTRALIA
Cape of
Good Hope
Argentina
Tasmania
Cape Horn
© 2014 Pearson Education, Inc.
Darwin in 1840, after
his return from the
voyage
New
Zealand
© 2014 Pearson Education, Inc.
Figure 21.5c
Figure 21.5d
The
Galápagos
Islands
Pinta
PACIFIC
OCEAN
Genovesa
Marchena
Santiago
Fernandina
Pinzón
Isabela
0
20 40
Kilometers
© 2014 Pearson Education, Inc.
Equator
Daphne
Islands
Santa Santa
Cruz
Fe
Florenza
San
Cristobal
HMS Beagle in port
Española
© 2014 Pearson Education, Inc.
12/7/2016
Video: Albatross Courtship Ritual
Video: Blue-footed Boobies Courtship Ritual
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Video: Galápagos Islands Overview
Video: Galápagos Marine Iguana
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Video: Galápagos Sea Lion
Video: Galápagos Tortoise
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Video: Soaring Hawk
Video: Snake Ritual Wrestling
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.6
Darwin’s Focus on Adaptation
 In reassessing his observations, Darwin perceived
adaptation to the environment and the origin of
new species as closely related processes
 From studies made years after Darwin’s voyage,
biologists have concluded that this is what
happened to the Galápagos finches
(a) Cactus-eater
(b) Insect-eater
(c) Seed-eater
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.6a
Figure 21.6b
(a) Cactus-eater
© 2014 Pearson Education, Inc.
(b) Insect-eater
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.6c
 In 1844, Darwin wrote an essay on natural selection
as the mechanism of descent with modification, but
did not introduce his theory publicly
 Natural selection is a process in which individuals with
favorable inherited traits are more likely to survive and
reproduce
(c) Seed-eater
 In June 1858, Darwin received a manuscript from
Alfred Russel Wallace, who had developed a theory of
natural selection similar to Darwin’s
 Darwin quickly finished The Origin of Species and
published it the next year
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
The Origin of Species
Descent with Modification
 Darwin explained three broad observations
 The unity of life
 The diversity of life
 The match between organisms and their
environment
© 2014 Pearson Education, Inc.
 Darwin never used the word evolution in the first
edition of The Origin of Species
 The phrase descent with modification summarized
Darwin’s perception of the unity of life
 The phrase refers to the view that all organisms
are related through descent from an ancestor that
lived in the remote past
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.7
 In the Darwinian view, the history of life is like a
tree with branches representing life’s diversity
 Darwin reasoned that large morphological gaps
between related groups could be explained by this
branching process and past extinction events
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.8
Figure 21.8a
Hyracoidea
(Hyraxes)
Hyracoidea
(Hyraxes)
Sirenia
(Manatees and relatives)
Sirenia
(Manatees and relatives)
†Moeritherium
†Barytherium
†Moeritherium
†Deinotherium
†Barytherium
†Mammut
†Platybelodon
†Deinotherium
†Stegodon
†Mammuthus
†Mammut
Elephas maximus (Asia)
Loxodonta africana (Africa)
Loxodonta cyclotis (Africa)
60
34
24
Millions of years ago
© 2014 Pearson Education, Inc.
5.5 2104 0
Years ago
60
34
24
Millions of years ago
© 2014 Pearson Education, Inc.
5.5 2104 0
Years ago
12/7/2016
Figure 21.8b
Artificial Selection, Natural Selection, and
Adaptation
†Platybelodon
 Darwin noted that humans have modified other
species by selecting and breeding individuals with
desired traits, a process called artificial selection
†Stegodon
†Mammuthus
 Darwin drew two inferences from two observations
Elephas maximus (Asia)
Loxodonta africana (Africa)
Loxodonta cyclotis (Africa)
60
34
24
5.5 2104 0
Millions of years ago
Years ago
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.9
Figure 21.9a
Cabbage
Selection
for apical
(tip) bud
Brussels
sprouts
Broccoli
Selection for
axillary (side)
buds
Selection for
flowers and stems
Selection
for stems
Selection
for leaves
Kale
© 2014 Pearson Education, Inc.
Wild mustard
Wild mustard
Kohlrabi
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.9b
Figure 21.9c
Brussels
sprouts
Kale
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.9d
Figure 21.9e
Cabbage
© 2014 Pearson Education, Inc.
Broccoli
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.9f
 Observation #1: Members of a population often
vary in their inherited traits
Kohlrabi
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.10
 Observation #2: All species can produce more
offspring than the environment can support, and
many of these offspring fail to survive and
reproduce
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.11
Spore
cloud
© 2014 Pearson Education, Inc.
 Inference #2: This unequal ability of individuals to
survive and reproduce will lead to the
accumulation of favorable traits in the population
over generations
 Inference #1: Individuals whose inherited traits
give them a higher probability of surviving and
reproducing in a given environment tend to leave
more offspring than other individuals
© 2014 Pearson Education, Inc.
 Darwin was influenced by Thomas Malthus, who
noted the potential for human population to
increase faster than food supplies and other
resources
 If some heritable traits are advantageous, these
will accumulate in a population over time, and this
will increase the frequency of individuals with
these traits
 This process explains the match between
organisms and their environment
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.12
Natural Selection: A Summary
 Individuals with certain heritable traits survive and
reproduce at a higher rate than other individuals
 Natural selection increases the match between
organisms and their environment over time
A flower mantid in Malaysia
A flower-eyed mantid in
South Africa
 If an environment changes over time, natural
selection may result in adaptation to these new
conditions and may give rise to new species
A leaf mantid in Borneo
© 2014 Pearson Education, Inc.
Figure 21.12a
© 2014 Pearson Education, Inc.
Figure 21.12b
A flower-eyed mantid in
South Africa
A flower mantid in Malaysia
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.12c
Video: Sea Horses
A leaf mantid in Borneo
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Concept 21.3: Evolution is supported by an
overwhelming amount of scientific evidence
 Note that individuals do not evolve; populations
evolve over time
 New discoveries continue to fill the gaps identified
by Darwin in The Origin of Species
 Natural selection can only increase or decrease
heritable traits that vary in a population
 There are four types of data that document the
pattern of evolution
 Adaptations vary with different environments
 Direct observations
 Homology
 The fossil record
 Biogeography
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Direct Observations of Evolutionary Change
 Two examples provide evidence for natural
selection: natural selection in response to
introduced plant species, and the evolution of
drug-resistant bacteria
© 2014 Pearson Education, Inc.
Natural Selection in Response to Introduced
Species
 Soapberry bugs use their “beak” to feed on seeds
within fruits
 Feeding is most effective when beak length is
closely matched to seed depth within the fruit
© 2014 Pearson Education, Inc.
 In southern Florida soapberry bugs feed on the
native balloon vine with larger fruit; they have
longer beaks
 In all cases, beak size has evolved in populations
that feed on introduced plants with fruits that are
smaller or larger than the native fruits
 In central Florida they feed on the introduced
goldenrain tree with smaller fruit; they have shorter
beaks
 These cases are examples of evolution by natural
selection
 Correlation between fruit size and beak size has
also been observed in Louisiana, Oklahoma, and
Australia
© 2014 Pearson Education, Inc.
 In Florida this evolution in beak size occurred in
less than 35 years
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.13
Figure 21.13a
Field Study
Results
10
Number of individuals
Field Study
Soapberry bug with beak
inserted in balloon vine
fruit
8
6
4
On native species,
balloon vine
(southern Florida)
Beak
2
0
Average for museum specimens
10
On introduced
species,
goldenrain tree
(central Florida)
8
6
4
2
0
6
7
8
9
Beak length (mm)
10
Soapberry bug with beak
inserted in balloon vine
fruit
11
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
The Evolution of Drug-Resistant Bacteria
 The bacterium Staphylococcus aureus is
commonly found on people
 Methicillin works by inhibiting a protein used by
bacteria to produce cell walls
 One strain, methicillin-resistant S. aureus (MRSA)
is a dangerous pathogen
 MRSA bacteria use a different protein in cell wall
production
 Resistance to penicillin evolved in S. aureus by
1945, two years after it was first widely used
 When exposed to methicillin, MRSA strains are
more likely to survive and reproduce than
nonresistant S. aureus strains
 Resistance to methicillin evolved in S. aureus by
1961, two years after it was first widely used
© 2014 Pearson Education, Inc.
 MRSA strains are now resistant to many
antibiotics
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.14
Figure 21.14a
2,750,000
1
250,000 base pairs
2,500,000
2,750,000
Chromosome map
of S. aureus clone USA300
1
2,500,000
Chromosome map
of S. aureus clone USA300
500,000
Key to adaptations
2,250,000
Methicillin resistance
Ability to colonize hosts
Increased disease severity
Increased gene exchange
(within species) and
toxin production
2,000,000
1,750,000
1,500,000
750,000
1,000,000
Annual hospital admissions
with MRSA (thousands)
250,000 base pairs
400
350
300
Key to adaptations
2,250,000
250
Methicillin resistance
Ability to colonize hosts
Increased disease severity
200
150
100
50
0
’93 ’94 ’95 ’96 ’97 ’98 ’99 ’00 ’01 ’02 ’03 ’04 ’05
Year
500,000
2,000,000
1,250,000
Increased gene exchange
(within species) and
toxin production
750,000
1,000,000
1,750,000
1,500,000
© 2014 Pearson Education, Inc.
1,250,000
© 2014 Pearson Education, Inc.
Figure 21.14b
Annual hospital admissions
with MRSA (thousands)
400
 Natural selection does not create new traits, but
edits or selects for traits already present in the
population
350
300
250
 The current, local environment determines which
traits will be selected for or selected against in any
specific population
200
150
100
50
0
© 2014 Pearson Education, Inc.
’93 ’94 ’95 ’96 ’97 ’98 ’99 ’00 ’01 ’02 ’03 ’04 ’05
Year
© 2014 Pearson Education, Inc.
12/7/2016
Homology
Anatomical and Molecular Homologies
 Homology is similarity resulting from common
ancestry
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 Homologous structures are anatomical
resemblances that represent variations on a
structural theme present in a common ancestor
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Figure 21.15
 Comparative embryology reveals anatomical
homologies not visible in adult organisms
Humerus
 For example, all vertebrate embryos have a postanal tail and pharyngeal arches
Radius
Ulna
Carpals
Metacarpals
Phalanges
Human
© 2014 Pearson Education, Inc.
Cat
Whale
Bat
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.16
Figure 21.16a
Pharyngeal
arches
Pharyngeal
arches
Post-anal
tail
Post-anal
tail
Chick embryo (LM)
Chick embryo (LM)
Human embryo
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.16b
 Vestigial structures are remnants of features that
served important functions in the organism’s
ancestors
Pharyngeal
arches
 Examples of homologies at the molecular level are
genes shared among organisms inherited from a
common ancestor
Post-anal
tail
Human embryo
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.17
Homologies and “Tree Thinking”
Branch point
Lungfishes
Mammals
2
Digit-bearing
limbs
Amnion
 Evolutionary trees can be made using different
types of data, for example, anatomical and DNA
sequence data
Lizards
and snakes
3
4
Homologous
characteristic
Crocodiles
Ostriches
6
Hawks and
other birds
© 2014 Pearson Education, Inc.
Figure 21.18
A Different Cause of Resemblance: Convergent
Evolution
 Convergent evolution is the evolution of similar,
or analogous, features in distantly related groups
 Analogous traits arise when groups independently
adapt to similar environments in similar ways
Sugar
glider
 Convergent evolution does not provide information
about ancestry
NORTH
AMERICA
AUSTRALIA
Flying squirrel
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Birds
5
Feathers
© 2014 Pearson Education, Inc.
Amniotes
 Homologies form nested patterns in evolutionary
trees
Amphibians
1
Tetrapods
 Evolutionary trees are hypotheses about the
relationships among different groups
12/7/2016
Figure 21.18a
Figure 21.18b
Sugar
glider
© 2014 Pearson Education, Inc.
Flying squirrel
© 2014 Pearson Education, Inc.
Figure 21.19
The Fossil Record
 The fossil record provides evidence of the
extinction of species, the origin of new groups, and
changes within groups over time
© 2014 Pearson Education, Inc.
Most mammals
Cetaceans and even-toed ungulates
(a) Canis (dog)
(b) Pakicetus
© 2014 Pearson Education, Inc.
(c) Sus (pig)
(d) Odocoileus (deer)
12/7/2016
Figure 21.19a
Figure 21.19b
(a) Canis (dog)
© 2014 Pearson Education, Inc.
(b) Pakicetus
© 2014 Pearson Education, Inc.
Figure 21.19c
Figure 21.19d
(c) Sus (pig)
© 2014 Pearson Education, Inc.
(d) Odocoileus (deer)
© 2014 Pearson Education, Inc.
12/7/2016
Figure 21.UN01
 Fossils can document important transitions
 For example, the transition from land to sea in the
ancestors of cetaceans
20 cm
Diacodexis, an early
even-toed ungulate
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.
Figure 21.20
Biogeography
Other even-toed
ungulates
 Biogeography, the scientific study of the
geographic distribution of species, provides
evidence of evolution
Hippopotamuses
†Pakicetus
 Earth’s continents were formerly united in a single
large continent called Pangaea, but have since
separated by continental drift
†Rodhocetus
†Dorudon
Common
ancestor
of cetaceans
0
50
40
30
60
Millions of years ago
© 2014 Pearson Education, Inc.
 An understanding of continent movement and
modern distribution of species allows us to predict
when and where different groups evolved
Living
cetaceans
Key
Pelvis
Femur
Tibia
Foot
© 2014 Pearson Education, Inc.
12/7/2016
What Is Theoretical About Darwin’s View of
Life?
 Endemic species are species that are not found
anywhere else in the world
 Islands have many endemic species that are often
closely related to species on the nearest mainland
or island
 Darwin explained that species from the mainland
colonized islands and gave rise to new species as
they adapted to new environments
© 2014 Pearson Education, Inc.
 In science, a theory accounts for many
observations and data and attempts to explain and
integrate a great variety of phenomena
 Darwin’s theory of evolution by natural selection
integrates diverse areas of biological study and
stimulates many new research questions
 Ongoing research adds to our understanding of
evolution
© 2014 Pearson Education, Inc.
Figure 21.UN02a
Figure 21.UN02b
Guppies
transplanted
Pools with
pike-cichlids
and guppies
© 2014 Pearson Education, Inc.
Pools with killifish,
but no guppies
prior to transplant
12
Area of colored
spots (mm2)
Number of
colored spots
12
10
8
6
4
2
0
Source
Transplanted
population population
© 2014 Pearson Education, Inc.
10
8
6
4
2
0
Source
Transplanted
population population
12/7/2016
Figure 21.UN03
Figure 21.UN04
Observations
Individuals in a population
vary in their heritable
characteristics.
Organisms produce more
offspring than the
environment can support.
Inferences
Individuals that are well suited
to their environment tend to leave more
offspring than other individuals.
and
Over time, favorable traits
accumulate in the population.
© 2014 Pearson Education, Inc.
Figure 21.UN05
© 2014 Pearson Education, Inc.
© 2014 Pearson Education, Inc.