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completely rewritten to incorporate new data and current thinking in terms of fungal phylogeny. The
present a more modern view of this critical topic. chapter was also edited for readability and clarity.
Chapter 17 The material on PCR was updated to include Chapter 26 Edits throughout emphasize the adaptations made by
reverse-transcription PCR (RT-PCR) and quantitative plants as they transitioned from an aquatic existence
RT-PCR. New sections were added to provide a more to a terrestrial one. The chapter was edited for clarity
contemporary perspective on molecular techniques and readability.
used in biotechnology. Modern techniques for Chapter 27 The chapter was edited throughout for clarity and
investigating gene function were added, including RNA readability.
interference, knockin mice, and the CRISPR genome
editing technology. Two new sections on modern Part VI Plant Form and Function
applications of biotechnology were added, which Chapter 29 Changes were made to section headings to emphasize
include the use of fluorescent in-situ hybridization and the key message of the section. The chapter was
gene chips in medical diagnostics. edited for clarity and readability.
Chapter 18 The section on genome sequencing was updated Chapter 30 The chapter was edited throughout for clarity and
to reflect current high-throughput technologies. The readability.
material on the Human Genome Project was updated Chapter 31 The chapter was edited throughout for clarity and
to include information about the Cancer Genome readability.
Project, and new material on the wheat genome
project reveals the challenges of sequencing larger, Part VII Animal Form and Function
repetitive genomes. The section on bioinformatics Chapter 32 The material on tissue types was rewritten to improve
was updated to include material on the use of mass readability, and a new figure was added showing
spectrometry in proteomics. the different types of epithelial tissue. The table
showing different types of neurons was changed to a
Part IV Evolution new figure to improve clarity. The material on animal
Chapter 19 The section on variation in natural populations was locomotion was edited for accuracy and clarity.
completely rewritten to take into account new data. Chapter 33 The material on supporting cells was updated
This includes extensive material on variation in to include new information. The description of
human populations. Learning objectives throughout membrane potential and action potentials was
were rewritten for consistency and clarity. rewritten to improve clarity and readability.
Chapter 20 The chapter was edited throughout for clarity and Chapter 34 The chapter throughout was edited for clarity and
consistency. The material on biogeography was readability.
updated and rewritten for clarity. Chapter 35 The material on distinctions between receptor types
Chapter 21 The chapters was edited throughout for increased was rewritten for clarity. A discussion of the ability
readability for students. The material on biological of steroid hormones to act via membrane receptors
species concept and speciation was rewritten to was added to update the material in this section.
improve clarity. The section on the action and control of G proteins
in signaling was updated and rewritten. Material on
Part V The Diversity of Life osmoregulation in fresh- and saltwater fish was added.
Chapter 22 The material on cladistics was edited for clarity Chapter 36 The material on blocks to polyspermy was updated
and readability. The material on the evolution of for clarity and accuracy. The section on embryonic
multicellularity was edited for clarity. stem cells and the the reversal of determination was
Chapter 23 The prokaryotic genetics section was rewritten extensively updated to include new data and provide
for clarity and to provide a modern perspective to context for the student.
complement the historical material. The material on
human diseases caused by infectious agents was Part VIII Ecology and Behavior
updated. A section on giant viruses was added to Chapter 37 The chapter was edited throughout for clarity and
provide information on this exciting new area. readability.
Chapter 24 The chapter was updated for accuracy, with new Chapter 38 The chapter was edited for clarity and accuracy. The
material added on control and treatment of malaria. data on human populations was updated.
Chapter 25 Updates focused on emphasizing the relationships Chapter 39 The material on mimicry was rewritten for clarity and
between structure and function in fungal forms. The accuracy. The material on keystone predator and
chapter was restructured, moving the more applied succession was updated for accuracy and readability.
aspects of mycology to the front of the chapter. Chapter 40 The chapter was edited throughout for clarity and
Changes to the Chytridiomycota section reflect readability.
changes to this edition vii

A Learning Path to Understanding Biology
Understanding Biology and its online assets have been carefully thought out and crafted to help students and professors work efficiently
and effectively through the material in the course, making the most of study time and instructional goals.
The Learning Path
of seeds introduces into the life cycle a dormant phase, which allows
Each chapter is ▶ 26.6 Seed Plants Were a Key Step the embryo to survive until environmental conditions are favorable
26
for further growth.
in Plant Evolution
organized in a way that A pollen grain is the male gametophyte
The history of the land plants is filled with evolutionary innovations
guides the studentCONCEPT 34.10 The Four Chambers of the Heart CONCEPT 34.11 The Circulatory Highway Isa few
Composed
Plants Seed plants produce two kinds of gametophytes—male and female—
allowing the ancestors of aquatic algae to colonize harsh and varied
each of which consists of just cells. Pollen grains, multicellular
terrestrial terrains. Early innovations made survival on land possible.
Contract in a Cycle of Arteries, Capillaries, and Veins
male gametophytes, are conveyed to the egg in the female gameto-
through the key Learni ng Path
Later innovations drove a radiation of plant life that continues to
phyte by wind or by a pollinator. In some seed plants, the sperm
change the landscape and the atmosphere, and that supports diverse moves toward the egg through a growing pollen tube. This elimi-
concepts.34.10.1 The Cardiac Cycle Drives the Cardiovascular 34.11.1 Arteries and Arterioles Have Evolved
for externalto
waterWithstand
animal life. nates the need through which sperm swim. In con-
Land Plants Are Multicellular
26.1
trast to the seedless plants, the whole male gametophyte, rather than
System High Pressures
Autotrophs Adapted to Terrestrial Life
Bryophytes Have a Dominant

26.2 The Seed Protects the Embryo just the sperm, moves to the female gametophyte.
Gametophyte Generation
A female gametophyte forms within the protection of the
34.10.2 Contraction of Heart Muscle Is Initiated by
Seedless Vascular Plants Evolved Roots,
26.3 34.11.2
LEARNING OBJECTIVE 26.6.1 Capillaries Form
List the evolutionary a Vast integuments,
advantages Network for Exchange
collectively forming the ovule. In angiosperms, the
▶
Stems, and Leaves
Every chapter opensAutorhythmic Cells Contraction is initiated by the SA

Lycophytes Have a Dominant
of seeds. ovules are completely enclosed within additional diploid sporophyte
tissue. The ovule and the surrounding protective tissue are called
34.11.3 Veins Have Less Muscle in Their Walls
26.4
Sporophyte Generation
with a Learning Pathnode, a natural pacemaker, and impulses then travel to thePterophytes Are Ferns and
Their Relatives
26.5
Seed-producing plants have come to dominate the terrestrial land-
scape over the last several hundred million years. Much of the remark-
the ovary. The ovary develops into the fruit.
AV node. Seed Plants Were a Key Step 34.11.4 The Lymphatic System Reclaims Fluids That
able success of seed plants, both gymnosperms and angiosperms, can
REVIEW OF CONCEPT 26.6 Leave
that walks through the
26.6
in Plant Evolution
be attributed to the evolution of the seed, an innovation that protects
Gymnosperms Are Plants
26.7
the Cardiovascular System
▲and provides food for delicate embryos. Seeds allow embryos to
A common ancestor that had seeds gave rise to the gymno-
with “Naked Seeds”
main concepts in the Angiosperms Are

Flowering Plants
26.8
remain dormant, germinating after a harsh winter or an extremely dry
Every concept is broken down intodispersal,
sectionsand can allow that cover
season has passed. Fruits, a later innovation, provided extra protection
for an extended
sperms and the angiosperms. Seeds protect the embryo, aid in
pause inskills or
the life cycle.
and enhanced the dispersal of embryos across a broader landscape.
Seed plants produce multicellular male and female gameto-
chapter. This helps ideas you should master. Learningphytes;
objectives at
Ancestral seed plants first appeared about 305 to 465 mya and
the beginning
the male gametophyte is a pollen grain, which is carried
preceded the appearance of gymnosperms and angiosperms. Seed
to the female gametophyte by wind or other means. The sperm
crospore. As the One sperm where
you understand unites with the egg and forms a zygote, which plants appear to have evolved from spore-bearing plants known as
of each section help you identify important concepts.
is within the pollen grain.
progymnosperms. Progymnosperms shared several features with
ature, the wall develops into an embryo sporophyte plant. The other sperm and ■
Why is water not essential for fertilization in seed plants?
the material fits in modern gymnosperms, including secondary vascular tissues (which
Assessing the Learning Path

The Connecting the Concepts
wn, leaving two the two polar nuclei unite, forming a triploid primary endo- allow for an increase in girth later in development). Some progymno-
feature at the end of each Part of ©
Dr
the text illustrates how five major .P
ar
concepts in biology relate to diverse v in
sperms had leaves. Their reproduction was very simple, and it is unclear
res have become the sperm
context of other
de
material in different chapters. This rS
et h
nucleus.
chapter contains material related to i
which particular group of progymnosperms gave rise to seed plants.
Gymnosperms Are Plants
the following concepts:
26.7
From an evolutionary and ecological perspective, the seed
concepts in theThe primary
chapter. endosperm nucleus begins dividing rapidly and
Structure Determines Function
Evolution Explains the Unity and

Diversity of Life
represents an important advance. The embryo is protected by an extra with “Naked Seeds”
layer or two of sporophyte tissue called the integument, creating the
omes beautifully repeatedly, becoming triploid endosperm tissue that may soon ovule (figure 26.16). Within the ovule, the megasporangium divides
act with others in consist of thousandsCONCEPT ▲34.1 Vertebrate
of cells. Endosperm Digestive
tissue can become an Systems Are CONCEPT 34.3 The Digestive Tract Is Regulated by
meiotically, producing a haploid megaspore. The megaspore produces
mas92413_ch26_570-593.indd 570 08/11/16 9:40 am
the egg that combines with the sperm, resulting in the zygote. Seeds
ale gametophyte Theextensive
chapter part of the Tubular
opening pages
seed Tracts
contain
in grasses such as icons
corn, that
and itlink to the
provides the Nervous System and Hormones
Ferns and Allies

also contain a food supply for the developing embryo.
Gymnosperms
Angiosperms
During development, the integuments harden to produce the
has areas called nutrients for the embryo in most flowering plants.
“Connecting the Concepts”
Understand feature that follow each Part in Understand seed coat. In addition to protecting the embryo from drought, the seed
can be easily dispersed. Perhaps even more significantly, the presence
emerge. Until recently, the nutritional,
1. The process oftriploid
digestionendosperm was of the following
occurs in which 1. The arrival of food into the stomach stimulates the release of
thebelieved
text. The Connecting
to be the ancestral the Concepts
state
structures show
ofintheangiosperms.
vertebrate how concepts
A recent
digestive system? (Select all that gastrin. What is the function of gastrin?
presented
analysis ofin different
extant, basal chapters
angiosperms
apply.) are related,
revealed connected
that diploid by a. Gastrin
Stored food
Integument
is a protease enzyme.
m its source (an endosperms were also common. a. Esophagus
The female gametophyte b. Smallinintestine b. Gastrin
(seed coat) activates pepsinogen.
unifying concepts in biology. c. eight,
These
Stomach
icons indicate what
d. Large
unifying There are four groups of living gymnosperms: conifero-
ring plant). Most these species has four, not nuclei. At the moment, it isintestine c. Gastrin
Embryo hydrolyzes proteins.
phytes, cycadophytes, gnetophytes, and ginkgophytes. They
ent plants and is concepts are illustrated

unclear whether diploid by the content in the
Applyor triploid endosperms are the morechapter. d. Gastrin stimulates the release of the
all lack pepsinogen and
flowers and fruits HCl. of angiosperms.
characteristic
In them all, the ovule, which becomes a seed, rests exposed
s, and other ani- primitive. 1. The primary function of the large intestine is to concentrate Apply on a scale (a modified shoot or leaf) and is not completely
enclosed by sporophyte tissues at the time of pollination.
rms, however, a wastes into solid form (feces) for release from the body. How 1. Eating a meal that contains 300 μm
a lot
Theofname
butter will trigger
gymnosperm literallywhich
means of“naked seed.”
Although the ovules are naked at the time of pollination, at
igma of its own does it accomplish this? the following?
Figure 26.16 Cross section of an ovule. maturity the seeds of gymnosperms are sometimes enclosed
Germination and growth a. Byof the additional
adding sporophyte cells from the mucosal layer a. Increased release of chyme byinto the duodenum
sporophyte tissues.
ay or may not be
© Biology Media/Science Source
tic compatibility b. By aabsorbing

As mentioned in section 26.6, seed may water
remain dormant for 582 Part V The Diversity b. Contraction
of Life of the gallbladder
many years, depending on c. By
thereleasing
species.salt
When environmental c. Inhibition of secretion
ma it has landed.
d. the
conditions become favorable, All seed
of theundergoes
above germination, and d. Increased secretion of pepsinogen and HCl
dense cytoplasm mas92413_ch26_570-593.indd 582 08/11/16 9:40 am
hrough an aper- the young sporophyteSynthesize

plant emerges. Again depending on the Synthesize
Chemical
species, the sporophyte1. may grow digestion involves
and develop for hydrolysis
many years reactions. Given what 1. Starving animals often exhibit swollen bodies rather than
sponds to chemi-
before becoming capable of you know about hydrolysis
reproduction, reactionsgrow
or it may quickly from chapter 3, write the emaciated ones in early stages of their deprivation. Why?
e embryo sac. It hydrolysis reaction to break down the disaccharide lactose.
nd grows down and produce flowers in a single growing season.
llen tube usually We present a more detailed description of reproduction in CONCEPT 34.4 Respiratory Systems Promote
icropyle, but the plants in chapter 30. CONCEPT 34.2 Food Is Processed As It Passes Efficient Exchange of Gases
is more rapid in
Through the Digestive Tract
Understand
Understand 1. Fick’s Law of Diffusion states the rate of diffusion is directly
erative cell, lags REVIEW OF CONCEPT
1. The 26.8
_______ and _______ play important roles in the proportional to
n the pollen tube, Angiosperms are characterizeddigestive
by process by producing
ovules that chemicals
at pollination are that are required to a. the area differences between the cross section of the blood
osses, ferns, and digest proteins, lipids, and carbohydrates. vessel and the tissue.
enclosed within an ovary at the base of a carpel, a structure
nts have no fla- a. liver; pancreas b. kidneys; appendix b. the pressure differences between the two sides of the
unique to the phylum; a fruit develops from the ovary. Evolution-
c. liver; gallbladder d. pancreas; gallbladder membrane and how large an area is available for diffusion.
e and sperm has ary innovations of angiosperms include flowers to attract pol-
Apply c. the pressure differences between the inside of the organism
linators, fruits to protect embryos and aid in their dispersal, and and the outside.
1. The small intestine is specialized for absorption because it
double fertilization, which provides endosperm to help nourish d. the temperature of the gas molecule.
on a. is the last section of the digestive tract and retains food the
the embryo. longest. Apply
troys a synergid ▲
■ What advantage doesb.anhas saclike extensions
angiosperm gain by along its length
producing a that collect food. 1. Which of the following evolutionary adaptations to increase
Both sperm are
fruit eaten by animals?c. has no outlet, so food remains within it for longer periods At the end gasof
exchange are matched
the chapter, eachcorrectly?
concept(Select
is all that apply.)at three
assessed
zation follows. of time. a. Beating cilia—increase surface area
d. has an extremely large surface area that allows extended different levels.
b. Bubbling gas into a fish tank—increases oxygenchapter prior to
On your first pass through the
▲ exposure to food. class, you will want to differences
concentration focus on questions at the “Understand”
SynthesizeReview of Concept questions c. Membranes lining the lungs are one-cell layer thick—
At the end of each section, level. As you gain greater
decreases distance
mastery, you can challenge yourself
1. How does the digestive system keep from being digested by
allow you to check yourthe understanding before moving
gastric secretions it produces? on with Applyd. and
A Synthesize
small diameter in questions that surface
bacteria—decreases require areahigher
to the next concept. cognitive skills.
Chapter 34 Fueling the Body’s Metabolism 819
viii
mas92413_ch34_789-821.indd 819 11/11/16 11:29 am
Think Like a Scientist
Scientific Thinking figures throughout Inquiry

◀ & Analysis
the text walk you through a scientific Why Do Diabetics Excrete
features at the end
Inquiry & Analysis

Effects of Eating on Blood Glucose Levels
Glucose in Their Urine?
experiment, laying out the Hypothesis, Late-onset diabetes is a serious and increasingly common
15
Diabetic
Normal
of every chapter take
disorder in which the body’s cells lose their ability to
Blood glucose level (mM )

Predictions, Test Procedures, Results, respond to insulin, a hormone that is needed to trigger
their uptake of glucose. As shown in the illustration, the 10 Kidney threshold
you into a scientific
binding of insulin to a receptor in the plasma membrane
and Conclusion. Some also challenge causes the rapid insertion of glucose transporter channels
into the plasma membrane, allowing the cell to take up
5
investigation in more
glucose. In diabetics, however, glucose molecules accu-
you to devise further experiments. mulate in the blood while the body’s cells starve for the
lack of them. In mild cases, blood glucose levels rise to detail, presenting you
several times the normal value of 4 mM; in severe 0
▼ untreated cases, blood glucose levels may become enor-

mously elevated, up to 25 times the normal value. A char-
0 1 2 3
Time (hours)
4 5 6
with experimental
acteristic symptom of even mild diabetes is the excretion
SCIENTIFIC THINKING
of dependent
The Cdc2 protein was the first identified cyclin- large amounts of glucose in the urine. The name of the
disorder,
kinase (Cdk), that is, a protein kinase enzyme that is diabetes mellitus, means “excessive secretion of
active only
results and challenging
sweet urine.” In normal individuals, by contrast, only long does it take for the normal person’s blood
Hypothesis: There are positive regulators of cell division.
Prediction: Frog oocytes are arrested in G2 of meiosis I. They can be

when complexed with cyclin. This finding led to the renaming
trace amounts of glucose are excreted. The kidney very
of MPF as mitosis-promoting factor, as its role was efficiently
clearly morereabsorbs glucose molecules from the fluid
glucose level to return to the level before the
test dose?
you to interpret the
general than simply promoting the maturation of through
frog
data. Associated online
induced to mature (undergo meiosis) by progesterone treatment. If passing it. Why doesn’t it do so in diabetic b. Comparing curves. Is the impact any different for
maturing oocytes contain a positive regulator of cell division, injection
oocytes. individuals? the diabetic person? How long does it take for the
of cytoplasm should induce an immature oocyte to undergo meiosis.
These Cdk enzymes are the key positive regulatorsThe of graph
the displays so-called glucose tolerance curves diabetic person’s blood glucose levels to return to
Test: Oocytes are induced with progesterone, then cytoplasm from
these maturing cells is injected into immature oocytes.
cell cycle, often called the engines that drive cell division.
more
After
Theperson (blue line) and a diabetic (red line).
for a normal
control of the cell cycle in higher eukaryotes is much com- without food, each individual drank a test
a night
the level before the test dose?
2. Interpreting Data
b. Is there any point at which the normal
activities can help you
plex than the simple single-cycle engine of yeast, dose of 100
but the grams of glucose dissolved in water. Blood glu-
yeast
Remove
cytoplasm
Inject
cytoplasm
model remains a useful framework for understanding
plex regulation.
cose levels
more were then monitored at 30-minute and 1-hour
com-
intervals. The dotted line indicates the kidney threshold,
individual’s blood glucose levels exceed the
kidney threshold? practice your data
the maximum concentration of blood glucose molecules b. Is there any point at which the diabetic individual’s
The Cell Cycle Can Be Halted

(about 10 mM) that the kidney is able to retrieve from the
fluid passing through it when all of its glucose-transporting
blood glucose levels do not exceed the kidney
threshold? analysis skills.
at Three Checkpoints channels are being utilized full-bore. 3. Making Inferences
a. Why do you suppose the diabetic individual
Progesterone- Arrested oocyte Oocyte in meiosis I LEARNING OBJECTIVE 10.6.2 Distinguish the roles of the three took so much longer to recover from the test
treated oocyte key checkpoints in the eukaryotic cell cycle. Analysis dose?
Result: Injected oocytes progress from G2 into meiosis I. b. Would you expect the normal individual to
Although for clarity we have divided the eukaryotic cell1.cycle
Applying
into Concepts excrete glucose? Explain. The diabetic individual?
Conclusion: The progesterone treatment causes production of a
arbitrary phases, the cell itself recognizes three phases, a. Reading
each a curve. What is the immediate impact Explain.
positive regulator of maturation: Maturation Promoting Factor (MPF).
marked by a checkpoint at which the cycle can be delayed on the
or normal individual’s blood glucose levels 4. Drawing Conclusions Why do diabetic individuals
of consuming the test dose of glucose? How secrete sweet urine?
Prediction: If mitosis is driven by positive regulators, then cytoplasm halted. The cell uses these three checkpoints both to assess its
from a mitotic cell should cause a G1 cell to enter mitosis. internal state and to integrate external signals (figure 10.16). The
Test: M phase cells are fused with G1 phase cells, then the nucleus checkpoints are located at the G1/S and G2/M boundaries, as well
from the G1 phase cell is monitored microscopically. as late in metaphase (the spindle checkpoint). Passage through Insulin not present Insulin present
these three checkpoints is controlled by the Cdk enzymes, as Glucose
follows. Insulin
The G1/S checkpoint

The G1/S checkpoint is the primary point at which the cell
M phase cell G1 phase cell Fused cells “decides” whether or not to divide. This checkpoint is therefore
the primary point at which external signals can influence events of
Conclusion: Cytoplasm from M phase cells contains a positive
the cycle. It is the phase during which growth factors affect the Insulin
regulator that causes a cell to enter mitosis. receptor Glucose
cycle, as well as the phase that links cell division to cell growth
transporter
Further Experiments: How can both of these experiments be and nutrition. channel Glucose
rationalized? What would be the next step in characterizing these

factors?
Chapter 34 Fueling the Body’s Metabolism 817
High
Figure 10.14 Discovery of positive regulator of cell MPF activity
Cyclin
division.
Concentration
mas92413_ch34_789-821.indd 817 11/11/16 11:29 am
necessary for cell-cycle progression. These studies indicated that

in yeast there were two critical control points: the commitment to
Low
Uncovering Connections Between Concepts

DNA synthesis (called START) and the commitment to mitosis.
One particular gene, named cdc2, was shown to be critical for G2 M G1 S G2 M G1 S G2 M
passing both of these boundaries.
MPF is cyclin plus cdc2 Figure 10.15 Correlation of MPF activity, amount of
All of these findings came together in an elegant fashion with the cyclin protein, and stages of the cell cycle. Cyclin
following three observations. First, the protein encoded by the
cdc2 gene was shown to be a protein kinase. Second, the purifica-
concentration and MPF activity are plotted vs. stage of the cell
cycle. Both change in a similar repeating pattern through the cell
A “Connecting
◀ the Concepts” feature at
tion and identification of MPF showed that it was composed of
both a cyclin component and a kinase component (figure 10.15).
cycle. The reason for this correlation is that cyclin is actually one
component of MPF, the other being a cyclin-dependent kinase
Connecting the Concepts Part II Biology of the Cell
the end of each Part in the text shows
Third, kinase was shown to be the Cdc2 protein! (Cdk). Together, these act as a positive regulator of cell division.
Life descended from early cells over 3.5 BYA. Over life’s history,
Chapter cells
10 have
Howadapted 211 into hundreds of different types. The diversity of
and diversified
Cells Divide
life is driven by how organisms acquire and process energy in the highly regulated reactions of photosynthesis and respiration. Plants have
how seemingly isolated concepts in
specialized organs, tissues, and cells to convert the Sun’s energy to chemical energy. Eukaryotes have specialized organelles that carry out the
regulated reactions of aerobic respiration. different chapters are connected by
Mason_ch10.indd 211
• Early life did not use oxygen.

08/09/16 5:14 pm
• Life descended from early cells that spontaneously

unifying concepts of biology.
arose over 3.5 billion years ago.
• Some prokaryotes adapted to anaerobic
environments by using molecules other than oxygen • All cells have a plasma membrane, cytoplasm, and DNA
as the final electron acceptor in cellular respiration. centrally located either in a nucleoid or a nucleus, but
vary in other structures.
• Fermentation is an anaerobic process that uses
organic compounds as electron acceptors. • Prokaryotic cells lack compartmentalization.
e o All • Plants cells have cell walls, chloroplasts, and large
Som isms 2
• Aerobic respiration uses oxygen as the final
are rgan central vacuoles, while animal cells lack these features.
electron acceptor and evolved after photosynthesis an O of o com isms
made oxygen available. org ’t use ne pos
n o e
do cell r mo d
s re
• Although chromosome structure differs
process of cell division.
• Leaves are the main organs for between prokaryotes and eukaryotes,
photosynthesis. Evolution DNA structure and function are similar.
photosynthesis
the controlled
are adapted
explains
arise from
• Leaf structure allows for gas exchange and • Prokaryotes have a single circular
All cells
Plants
minimizes water loss, and cells within contain chromosome and divide clonally by
for
chloroplasts.
the unity and binary fission. Eukaryotic division is
• Pigment molecules in plants absorb specific diversity complex, due to genome size.
wavelengths of light, with different pigment of life • Larger and more complex genomes
molecules expanding the spectrum of the required the evolution of mitosis to
Sun’s energy plants absorb. segregate chromosomes accurately.
• Plants use rubisco to fix CO2 but rubisco also
binds oxygen, reversing carbon fixation.
• C4 and CAM plants have adaptations to
minimize photorespiration.
• Photosynthetic organisms
capture the Sun’s energy
The ycle y
and use this energy to

is c ulat
c efull
reg
power all life.

cell
ar ed
organic molecules
• Light-dependent reactions
energy to make
make ATP and NADPH Living systems Living systems

uses light
synthesis
Photo-
which power the synthesis

of organic molecules from transform depend on
atmospheric CO2 in the energy & matter information
res cept ase
light-independent
transactions
re in kin
ph pond ors
reactions.
to
ion -
lat hory
Th ize
te
• Photosystems are erm

oxidic
Pro
osp
important structures that

go odyn lls
Ce rgan nds
capture light energy and ve a Critical
en rn a mic o ou
p
convert it into chemical cha ergy ll s m
co driv sm
e processes
energy in the form of ng to boli are regulated
es ta
excited electrons. me by feedback
inhibition
• The biological world is an open system in Now that you’ve seen two
which energy flows from the Sun and is • Oxidation-reduction (redox) reactions capture
energy from food in the form of ATP. examples of Connecting the
eventually lost as heat. Concepts, fill in the supporting
• All of life’s activities involve changes in energy • Electron carriers, like NAD+, shuttle electrons details for “Living systems
that follow the laws of thermodynamics. and their energy from one molecule to depend on information
another. transactions” using the
• Photosynthesis and cellular respiration are
energy-transforming reactions critical to the • Energy released during cellular respiration is concepts provided.
living world. used to create proton gradients that drive
ATP synthesis.
a learning path to understanding biolog y ix

Using Connect® and Understanding Biology, 2nd Edition
Prepare for the Course

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the course working knowledge of basic chemistry and cellular biology. For this reason,
many biology students struggle the first few weeks of class. LearnSmart Prep
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Professors can assign modules in LearnSmart Prep to help students get up
to speed on core concepts, or students can access LearnSmart Prep directly
through the LearnSmart Prep link.
Prepare before
class LearnSmart Prep is an adaptive
learning tool designed to increase
student success and aid retention
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biology students can master some of the most fundamental and
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Tying it all
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2 As you work through the learning plan, the program

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(e.g., animation or tutorial) on that concept, then will
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Acknowledgments
A revision of this scope relies on the talents and efforts of many digital team, whom we owe a great deal for their efforts to help
people working behind the scenes and we have benefited us move toward the future.
greatly from their assistance.
The copy editor, Deb DeBord, labored many hours and Digital Team: Ian Quitadamo, Central Washington University,
always improved the clarity and consistency of the text. serves as the Lead Digital Author in Majors Biology, directing
She has made a tremendous contribution to the quality the development plan for the digital content. Scott Cooper,
of the final product. We were fortunate to work with MPS University of Wisconsin–LaCrosse, serves as the Lead Connect
Limited to update the art program and improve the layout of digital content team, and Michelle Pass, University of North
the pages. Our close collaboration resulted in a text that is Carolina–Charlotte, serves as the Lead for the SmartBook
pedagogically effective as well as more beautiful than any development team.
other biology text on the market. Throughout this edition we have had the support of spouses
We have the continued support of an excellent team at McGraw- and children, who have seen less of us than they might
Hill Education. Justin Wyatt, the Brand Manager for Biology, have liked because of the pressures of getting this revision
has been a steady leader during a time of change. The Lead completed. They have adapted to the many hours this book
Product Developer, Liz Sievers, provided support in so many draws us away from them, and, even more than us, looked
ways it would be impossible to name them all. April Southwood, forward to its completion.
Lead Content Project Manager, and Matt Backhous, Designer, In the end, the people we owe the most are the generations of
ensured our text was elegantly designed. Patrick Reidy, students who have passed through our lecture halls. They have
Marketing Manager, is always a sounding board for more than taught us at least as much as we have taught them, and their
just marketing, and many more people behind the scenes have questions and suggestions continue to improve the text and
all contributed to the success of our text. This includes the supplementary materials.
xvii
Contents
About the Authors iv
PART II Biology
Changes to This Edition vi
of the Cell 66
A Learning Path to Understanding Biology viii
Acknowledgments xvii 4 Cell Structure 66
Concept 4.1 All Living Organisms Are Composed of Cells 67
Concept 4.2 Prokaryotic Cells Lack Interior Organization 70
PART I The Molecular Concept 4.3

Concept 4.4
Eukaryotic Cells Are Highly Compartmentalized 72
Membranes Organize the Cell Interior into Functional
Basis of Life 1 Compartments 76
Concept 4.5 Mitochondria and Chloroplasts Are Energy-Processing
Organelles 80
1 The Science of Biology 1
Concept 4.6 An Internal Skeleton Supports the Shape of Cells 82
Concept 1.1 The Diversity of Life Is Overwhelming 2 Concept 4.7 Extracellular Structures Protect Cells 85
Concept 1.2 Biology Is the Science of Life 3 Concept 4.8 Cell-to-Cell Connections Determine How Adjacent
Concept 1.3 Science Is Based on Both Observation and Cells Interact 86
Reasoning 6 Inquiry & Analysis 90
Concept 1.4 The Study of Evolution Is a Good Example Retracing the Learning Path 91
of Scientific Inquiry 9
Assessing the Learning Path 92
Concept 1.5 A Few Important Concepts Form the Core of
Biology 13
5 Membranes 94
Inquiry & Analysis 17
Retracing the Learning Path 18 Concept 5.1 Membranes Are Phospholipid Bilayers with Embedded
Proteins 95
Concept 5.2 Phospholipids Provide a Membrane’s Structural
Foundation 98
2 The Nature of Molecules and the Properties Concept 5.3 Membrane Proteins Enable a Broad Range of
of Water 21 Interactions with the Environment 99
Concept 5.4 Passive Transport Moves Molecules Across
Concept 2.1 All Matter Is Composed of Atoms 22
Membranes by Diffusion 101
Concept 2.2 The Elements in Living Systems Have Low Atomic
Concept 5.5 Active Transport Across Membranes Requires
Masses 25
Energy 105
Concept 2.3 Molecules Are Collections of Atoms Held Together by
Concept 5.6 Bulky Materials Cross Membranes Within
Chemical Bonds 27
Vesicles 108
Concept 2.4 The Properties of Water Result from Its
Polar Nature 30
Retracing the Learning Path 112
Concept 2.5 Water Molecules Can Dissociate into Ions 34
6 Energy and Metabolism 115
Concept 6.1 Energy Flows Through Living Systems 116
3 The Chemical Building Blocks of Life 40 Concept 6.2 The Laws of Thermodynamics Govern All Energy
Changes 117
Concept 3.1 Carbon Provides the Framework of Biological
Concept 6.3 ATP Is the Energy Currency of Cells 119
Molecules 41
Concept 6.4 Enzymes Speed Up Reactions by Lowering Activation
Concept 3.2 Carbohydrates Form Both Structural
Energy 121
and Energy-Storing Molecules 43
Concept 6.5 Metabolism Is the Sum of a Cell’s Chemical
Concept 3.3 Proteins Are the Tools of the Cell 47
Activities 125
Concept 3.4 Nucleic Acids Store and Express Genetic
Information 55
Concept 3.5 Hydrophobic Lipids Form Fats and Membranes 58
xviii
7 How Cells Harvest Energy 131 10 How Cells Divide 198
Concept 7.1 Cells Harvest Energy from Organic Compounds by Concept 10.1 Bacterial Cell Division Is Clonal 199
Oxidation 132 Concept 10.2 Eukaryotes Have Large, Linear Chromosomes 200
Concept 7.2 Glycolysis Splits Glucose and Yields a Small Amount Concept 10.3 The Eukaryotic Cell Cycle Is Complex and Highly
of ATP 136 Organized 203
Concept 7.3 The Krebs Cycle Is the Oxidative Core of Cellular Concept 10.4 During Interphase, Cells Grow and Prepare for
Respiration 139 Mitosis 204
Concept 7.4 Electrons Harvested by Oxidation Pass Along an Concept 10.5 In Mitosis, Chromosomes Segregate 206
Electron Transport Chain 142
Concept 10.6 Events of the Cell Cycle Are Carefully
Concept 7.5 The Total Energy Yield of Aerobic Respiration Far Regulated 210
Exceeds That of Glycolysis 146
Concept 10.7 Cancer Is a Failure of Cell-Cycle Control 214
Concept 7.6 Aerobic Respiration Is Regulated by Feedback
Inhibition 147 Inquiry & Analysis 217
Concept 7.7 Some Forms of Energy Metabolism Do Not Require Retracing the Learning Path 218
O2 148 Assessing the Learning Path 219
Concept 7.8 Carbohydrates Are Not the Only Energy Source Used
by Heterotrophs 149
Retracing the Learning Path 153 PART III Genetic and
Assessing the Learning Path 154 Molecular Biology 222
8 Photosynthesis 156 11 Sexual Reproduction and Meiosis 222
Concept 8.1 Photosynthesis Uses Sunlight to Power the Synthesis
of Organic Molecules 157 Concept 11.1 Sexual Reproduction Requires Meiosis 223
Concept 8.2 Experiments Revealed That Photosynthesis Is a Concept 11.2 Meiosis Consists of Two Divisions with One Round
Chemical Process 159 of DNA Replication 224
Concept 8.3 Pigments Capture Energy from Sunlight 160 Concept 11.3 The Process of Meiosis Involves Intimate Interactions
Between Homologues 225
Concept 8.4 Photosynthetic Pigments Are Organized into
Photosystems 163 Concept 11.4 Meiosis Has Four Distinct Features 230
Concept 8.5 Energy from Sunlight Is Used to Produce a Proton Concept 11.5 Genetic Variation Is the Evolutionary Consequence
Gradient 165 of Sex 232
Concept 8.6 Using ATP and NADPH from the Light Reactions, Inquiry & Analysis 234
CO2 Is Incorporated into Organic Molecules 170 Retracing the Learning Path 235
Concept 8.7 Photorespiration Short-Circuits Photosynthesis 172 Assessing the Learning Path 236
Retracing the Learning Path 176 12 Patterns of Inheritance 238
Assessing the Learning Path 177 Concept 12.1 Experiments Carried Out by Mendel Explain
Heredity 239
9 Cell Communication 179 Concept 12.2 Mendel’s Principle of Segregation Accounts for 3:1
Phenotypic Ratios 241
Concept 9.1 The Cells of Multicellular Organisms
Communicate 180 Concept 12.3 Mendel’s Principle of Independent Assortment Asserts
That Genes Segregate Independently 245
Concept 9.2 Signal Transduction Begins with Cellular
Receptors 182 Concept 12.4 Probability Allows Us to Predict the Results of
Crosses 246
Concept 9.3 Intracellular Receptors Respond to Signals by Regulating
Gene Expression 184 Concept 12.5 Genotype Dictates Phenotype by Specifying Protein
Sequences 248
Concept 9.4 Protein Kinase Receptors Respond to Signals by
Phosphorylating Proteins 185 Concept 12.6 Extending Mendel’s Model Provides a Clearer View of
Genetics in Action 249
Concept 9.5 G Protein–Coupled Receptors Respond to Signals
Through Effector Proteins 189 Inquiry & Analysis 255
Inquiry & Analysis 194 Retracing the Learning Path 256
Retracing the Learning Path 195 Assessing the Learning Path 257
contents xix
13 The Chromosomal Basis 16 Control of Gene Expression 332
of Inheritance 259 Concept 16.1 All Organisms Control Expression of Their
Genes 333
Concept 13.1 Chromosomes Are the Vehicles of Mendelian
Inheritance 260 Concept 16.2 Regulatory Proteins Control Genes by Interacting with
Specific DNA Nucleotide Sequences 334
Concept 13.2 Some Genes Do Not Assort Independently:
Linkage 263 Concept 16.3 Prokaryotes Regulate Their Genes in Clusters 336
Concept 13.3 Genetic Crosses Provide Data for Genetic Maps 265 Concept 16.4 Transcription Factors Control Gene Transcription in
Eukaryotes 340
Concept 13.4 Changes in Chromosome Number Can Have Drastic
Effects 267 Concept 16.5 Chromatin Structure Affects Gene Expression 344
Concept 13.5 Inheritance in Humans Can Be Studied by Analyzing Concept 16.6 Eukaryotic Genes Are Also Regulated After
Pedigrees 268 Transcription 345
Concept 13.6 There Are Two Major Exceptions to Chromosomal Concept 16.7 Gene Regulation Determines How Cells Will
Inheritance 273 Develop 351
Inquiry & Analysis 276 Inquiry & Analysis 354
Retracing the Learning Path 277 Retracing the Learning Path 355
Assessing the Learning Path 278 Assessing the Learning Path 356
14 DNA: The Genetic Material 280 17 Biotechnology 358

Concept 14.1 DNA Is the Genetic Material 281 Concept 17.1 Enzymes Allow the Creation of Recombinant
Molecules in Vitro 359
Concept 14.2 The DNA Molecule Is a Double Helix 283
Concept 17.2 The Polymerase Chain Reaction Is Used to Amplify
Concept 14.3 Both Strands Are Copied During DNA Specific DNA Sequences 363
Replication 287
Concept 17.3 Molecular Tools Allow Us to Analyze and Modify
Concept 14.4 Prokaryotes Organize the Enzymes Used to Duplicate Genetic Variation 366
DNA 290
Concept 17.4 Transgenic Organisms Are Used to Analyze Gene
Concept 14.5 Eukaryotic Chromosomes Are Large and Linear 295 Function 368
Concept 14.6 Cells Repair Damaged DNA 297 Concept 17.5 Genetic Tools Are Changing Modern Medicine 371
Inquiry & Analysis 300 Concept 17.6 Genetic Engineering Is Used in Industry and
Retracing the Learning Path 301 Agriculture 374
Assessing the Learning Path 302 Inquiry & Analysis 378
15 Genes and How They Work 304 Assessing the Learning Path 380
Concept 15.1 Experiments Have Revealed the Nature of Genes 305
Concept 15.2 The Genetic Code Relates Information in DNA and 18 Genomics 383
Protein 307 Concept 18.1 Mapping Identifies and Locates Functional Elements
Concept 15.3 Prokaryotes Exhibit All the Basic Features of in Genomes 384
Transcription 310 Concept 18.2 The Modernization of DNA Sequencing Has
Concept 15.4 Eukaryotes Use Three Polymerases, and Extensively Accelerated Discovery 387
Modify Transcripts 313 Concept 18.3 Genome Projects Reveal Insights into Medicine and
Concept 15.5 Eukaryotic Genes May Contain Noncoding Agriculture 390
Sequences 315 Concept 18.4 Genome Annotation Assigns Functional Information to
Concept 15.6 The Ribosome Is the Machine of Protein Genomes 392
Synthesis 317 Concept 18.5 Genome Comparisons Provide Information About
Concept 15.7 The Process of Translation Is Complex and Energy- Genomic Structure and Function 395
Expensive 320 Concept 18.6 Comparative Genomics Informs Evolutionary
Concept 15.8 Mutations Are Alterations in the Sequence, Number, or Biology 400
Position of Genes 325 Inquiry & Analysis 404
xx contents
PART IV PART V The Diversity
Evolution 409 of Life 477
19 Genes Within Populations 409 22 Systematics and Phylogeny 477
Concept 19.1 Natural Populations Exhibit Genetic Variation 410 Concept 22.1 Systematics Reconstructs Evolutionary
Concept 19.2 Frequencies of Alleles Can Change 412 Relationships 478
Concept 19.3 Five Agents Are Responsible for Evolutionary Concept 22.2 Cladistics Focuses on Traits Derived from a Common
Change 414 Ancestor 479
Concept 19.4 Selection Can Act on Traits Affected by Many Concept 22.3 Classification Is a Labeling Process, Not an
Genes 419 Evolutionary Reconstruction 483
Concept 19.5 Natural Selection Can Be Studied Experimentally 421 Concept 22.4 Taxonomy Attempts to Classify Organisms in an
Evolutionary Context 486
Concept 19.6 Fitness Is a Measure of Evolutionary Success 423
Concept 22.5 The Largest Taxons Are Domains 488
Concept 19.7 Interacting Evolutionary Forces Maintain Variation 423
23 Prokaryotes and Viruses 497

20 The Evidence for Evolution 431
Concept 23.1 Prokaryotes Are the Most Ancient Organisms 498
Concept 20.1 The Beaks of Darwin’s Finches Provide Evidence of
Natural Selection 432 Concept 23.2 Prokaryotes Have an Organized but Simple
Structure 501
Concept 20.2 Peppered Moths and Industrial Melanism Illustrate
Natural Selection in Action 435 Concept 23.3 The Genetics of Prokaryotes Focuses on DNA
Transfer 505
Concept 20.3 Human-Initiated Artificial Selection Is Also a Powerful
Agent of Change 437 Concept 23.4 Prokaryotic Metabolism Is Quite Diverse 509
Concept 20.4 Fossils Provide Direct Evidence of Evolution 439 Concept 23.5 Bacteria Cause Important Human Diseases 510
Concept 20.5 Anatomical Evidence for Evolution Is Extensive Concept 23.6 Viruses Are Not Organisms 511
and Persuasive 442 Concept 23.7 Bacterial Viruses Infect by DNA Injection 514
Concept 20.6 Genes Carry a Molecular Record of the Evolutionary Concept 23.8 Animal Viruses Infect by Endocytosis 515
Past 445 Inquiry & Analysis 518
Concept 20.7 Natural Selection Favors Convergent Evolution in Retracing the Learning Path 519
Similar Environments 446
Concept 20.8 Addressing Common Criticisms of Evolutionary
Theory 448
24 Protists 522
Retracing the Learning Path 451 Concept 24.1 Protists, the First Eukaryotes, Arose by
Endosymbiosis 523
Concept 24.2 Biologically, Protists Are a Very Diverse
Group 525
21 The Origin of Species 454 Concept 24.3 The Rough Outlines of Protist Phylogeny Are
Concept 21.1 The Biological Species Concept Highlights Becoming Clearer 527
Reproductive Isolation 455 Concept 24.4 Excavata Are Flagellated Protists Lacking
Concept 21.2 Natural Selection May Reinforce Reproductive Mitochondria 528
Isolation 460 Concept 24.5 Chromalveolata Seem to Have Originated by
Concept 21.3 Natural Selection and Genetic Drift Play Key Roles in Secondary Symbiosis 531
Speciation 461 Concept 24.6 Rhizaria Have Silicon Exoskeletons or Limestone
Concept 21.4 Speciation Is Influenced by Geography 462 Shells 537
Concept 21.5 Adaptive Radiation Requires Both Speciation and Concept 24.7 Archaeplastida Are Descended from a Single
Habitat Diversity 464 Endosymbiosis Event 538
Concept 21.6 The Pace of Evolution Varies 469 Concept 24.8 Amoebozoa and Opisthokonta Are Closely
Concept 21.7 Speciation and Extinction Have Molded Biodiversity Related 541
Through Time 470 Inquiry & Analysis 544
contents xxi
25 Fungi 548 28 Vertebrates 625
Concept 25.1 Fungi Have Unique Reproductive and Nutritional Concept 28.1 Nonvertebrate Chordates Do Not Form
Strategies 549 Bone 626
Concept 25.2 Fungi Have an Enormous Ecological Impact 552 Concept 28.2 Almost All Chordates Are Vertebrates 627
Concept 25.3 Fungi Are Important Plant and Animal Pathogens 555 Concept 28.3 Fishes Are the Earliest and Most Diverse
Concept 25.4 Fungi Are Taxonomically Diverse 556 Vertebrates 628
Concept 25.5 Microsporidia Are Unicellular Parasites 557 Concept 28.4 Amphibians Were the First Terrestrial
Vertebrates 632
Concept 25.6 Chytridiomycota and Relatives: Fungi with
Zoospores 558 Concept 28.5 Reptiles Are Fully Adapted to Terrestrial
Living 634
Concept 25.7 Zygomycota Produce Zygotes 559
Concept 28.6 Birds Are Essentially Flying Reptiles 638
Concept 25.8 Glomeromycota Are Asexual Plant Symbionts 560
Concept 28.7 Mammals Are the Least Diverse of
Concept 25.9 Basidiomycota Are the Mushroom Fungi 561 Vertebrates 641
Concept 25.10 Ascomycota Are the Most Diverse Phylum of Fungi 562 Concept 28.8 Primates Include Anthropoids and
Inquiry & Analysis 565 Prosimians 644
Retracing the Learning Path 566 Inquiry & Analysis 651
Assessing the Learning Path 567 Retracing the Learning Path 652
26 Plants 570
Concept 26.1 Land Plants Are Multicellular Autotrophs Adapted to
Terrestrial Life 571
Concept 26.2 Bryophytes Have a Dominant Gametophyte PART VI Plant Form
Generation 573
Concept 26.3 Seedless Vascular Plants Evolved Roots, Stems,
and Function 656
and Leaves 576
Concept 26.4 Lycophytes Have a Dominant Sporophyte 29 Plant Form 656
Generation 577 Concept 29.1 Meristems Articulate the Plant Body 657
Concept 26.5 Pterophytes Are Ferns and Their Relatives 578 Concept 29.2 Plants Contain Three Principal Tissues 660
Concept 26.6 Seed Plants Were a Key Step in Plant Evolution 582 Concept 29.3 Roots Have Four Growth Zones 665
Concept 26.7 Gymnosperms Are Plants with “Naked Seeds” 582 Concept 29.4 Stems Provide Support for Aboveground
Concept 26.8 Angiosperms Are Flowering Plants 586 Organs 670
Inquiry & Analysis 590 Concept 29.5 Leaves Are a Plant’s Photosynthetic Organs 674
27 Animal Diversity 594
Concept 27.1 The Diversity of Animal Body Plans Arose by a Series 30 Flowering Plant Reproduction 681
of Evolutionary Innovations 595 Concept 30.1 Reproduction Starts with Flowering 682
Concept 27.2 Molecular Data Are Clarifying the Animal Concept 30.2 Flowers Attract Pollinators 684
Phylogenetic Tree 600
Concept 30.3 Fertilization Leads to Embryogenesis 689
Concept 27.3 True Tissue Evolved in Simple Animals 604
Concept 30.4 Seeds Protect Angiosperm Embryos 692
Concept 27.4 Platyzoans Are Very Simple Bilaterians 606
Concept 30.5 Fruits Promote Seed Dispersal 694
Concept 27.5 Mollusks and Annelids Are the Largest Groups of
Lophotrochozoans 608 Concept 30.6 Germination Begins Seedling Growth 696
Concept 27.6 Lophophorates Are Very Simple Marine Concept 30.7 Plant Life Spans Vary Widely 698
Organisms 612 Concept 30.8 Asexual Reproduction Is Common Among
Concept 27.7 Nematodes and Arthropods Are Both Large Groups Flowering Plants 699
of Ecdysozoans 613 Inquiry & Analysis 701
Concept 27.8 Deuterostomes Are Composed of Echinoderms and Retracing the Learning Path 702
Chordates 618 Assessing the Learning Path 703
xxii contents
31 The Living Plant 706 Concept 33.9 Sounds and Body Position Are Sensed by Vibration
Detectors 776
Concept 31.1 Water Moves Through Plants Based on Potential Concept 33.10 Taste, Smell, and pH Senses Utilize
Differences 707 Chemoreceptors 779
Concept 31.2 Roots Absorb Minerals and Water 710 Concept 33.11 Vision Employs Photoreceptors to Perceive Objects
Concept 31.3 Xylem Transports Water from Root to Shoot 711 at a Distance 780
Concept 31.4 Transpiration Rate Reflects Environmental Inquiry & Analysis 785
Conditions 714 Retracing the Learning Path 786
Concept 31.5 Plants Are Adapted to Water Stress 716 Assessing the Learning Path 787
Concept 31.6 Phloem Transports Organic Molecules 717
Concept 31.7 Plants Require a Variety of Nutrients 719 34 Fueling the Body’s Metabolism 789
Concept 31.8 Plants Use Hormones to Regulate Growth 720
Concept 34.1 Vertebrate Digestive Systems Are Tubular Tracts 790
Concept 31.9 Plant Growth Is Responsive to Light 725
Concept 34.2 Food Is Processed As It Passes Through the Digestive
Concept 31.10 Plant Growth Is Sensitive to Gravity 728 Tract 791
Inquiry & Analysis 731 Concept 34.3 The Digestive Tract Is Regulated by the Nervous
Retracing the Learning Path 732 System and Hormones 796
Assessing the Learning Path 733 Concept 34.4 Respiratory Systems Promote Efficient Exchange
of Gases 797
Concept 34.5 Gills Provide for Efficient Gas Exchange in Water 798
Concept 34.6 Lungs Are the Respiratory Organs of Terrestrial
PART VII Animal Form Vertebrates 800
Concept 34.7 Oxygen and Carbon Dioxide Are Transported by
and Function 736 Fundamentally Different Mechanisms 804
Concept 34.8 Circulating Blood Carries Metabolites and Gases to
32 The Animal Body and How the Tissues 806
Concept 34.9 Vertebrate Circulatory Systems Put a Premium on
It Moves 736 Efficient Circulation 808
Concept 32.1 The Vertebrate Body Has a Hierarchical Concept 34.10 The Four Chambers of the Heart Contract in a
Organization 737 Cycle 811
Concept 32.2 Epithelial Tissue Forms Barriers Within the Body 739 Concept 34.11 The Circulatory Highway Is Composed of Arteries,
Concept 32.3 Nerve Tissue Conducts Signals Rapidly 741 Capillaries, and Veins 813
Concept 32.4 Connective Tissue Supports the Body 742 Inquiry & Analysis 817
Concept 32.5 Muscle Tissue Powers the Body’s Movements 743 Retracing the Learning Path 818
Concept 32.6 Skeletal Systems Anchor the Body’s Muscles 745 Assessing the Learning Path 819
Concept 32.7 Vertebrate Endoskeletons Are Made of Bone 746
Concept 32.8 Muscles Contract Because Their Myofilaments 35 Maintaining Homeostasis 822
Shorten 750 Concept 35.1 Homeostasis Maintains a Constant Internal
Concept 32.9 Animal Locomotion Takes Many Forms 754 Environment 823
Inquiry & Analysis 757 Concept 35.2 Hormones Are Chemical Messages That Direct Body
Retracing the Learning Path 758 Processes 827
Assessing the Learning Path 759 Concept 35.3 The Pituitary and the Hypothalamus Are the Body’s
Control Centers 831
33 The Nervous System 761 Concept 35.4 Peripheral Endocrine Glands Play Major Roles in
Homeostasis 834
Concept 33.1 The Nervous System Directs the Body’s Actions 762 Concept 35.5 Animals Are Osmoconformers or Osmoregulators 838
Concept 33.2 Neurons Maintain a Resting Potential Across the Concept 35.6 The Kidney Maintains Osmotic Homeostasis in
Plasma Membrane 763 Mammals 839
Concept 33.3 Action Potentials Propagate Nerve Impulses 765 Concept 35.7 Hormones Control Osmoregulation 844
Concept 33.4 Synapses Are Where Neurons Communicate with Concept 35.8 The Immune System Defends the Body 845
Other Cells 767
Concept 35.9 Cell-Mediated Immunity Involves Helper and Killer
Concept 33.5 The Central Nervous System Includes the Brain and T Cells 850
Spinal Cord 770
Concept 35.10 In Humoral Immunity, B Cells Produce Protective
Concept 33.6 The Peripheral Nervous System Consists of Both Antibodies 852
Sensory and Motor Neurons 773
Concept 33.7 Sensory Receptors Provide Information About the
Body’s Environment 774 Retracing the Learning Path 857
Concept 33.8 Mechanoreceptors Sense Touch and Pressure 775 Assessing the Learning Path 858
contents xxiii
36 Reproduction and Development 860 Concept 38.3 Evolution Favors Life Histories That Maximize
Lifetime Reproductive Success 925
Concept 36.1 Mammals Are Viviparous 861 Concept 38.4 Environment Limits Population Growth 927
Concept 36.2 The Human Male Reproductive System Is Typical Concept 38.5 Resource Availability Regulates Population
of Mammals 862 Growth 929
Concept 36.3 The Human Female Reproductive System Undergoes Concept 38.6 Earth’s Human Population Is Growing
Cyclic Gamete Development 865 Explosively 932
Concept 36.4 The First Step in Development Is Fertilization 870 Inquiry & Analysis 936
Concept 36.5 Cells of the Early Embryo Are Totipotent 873 Retracing the Learning Path 937
Concept 36.6 Cleavage Leads to the Blastula Stage 878 Assessing the Learning Path 938
Concept 36.7 Gastrulation Forms the Basic Body Plan of the
Embryo 879
39 Community Ecology 940
Concept 36.8 The Body’s Organs Form in Organogenesis 881
Concept 36.9 Human Development Takes Nine Months 884 Concept 39.1 Competition Shapes How Species Live Together in
Communities 941
Concept 39.2 Predator–Prey Relationships Foster Coevolution 945
Concept 39.3 Cooperation Among Species Can Lead to
Assessing the Learning Path 890 Coevolution 949
Concept 39.4 Ecological Succession Is a Consequence of Habitat
Alteration 954
Concept 39.5 Chemical Elements Move Through Ecosystems in
PART VIII Ecology Biogeochemical Cycles 956
and Behavior 893 Concept 39.6 Energy Flows Through Ecosystems in One
Direction 961
Concept 39.7 Biodiversity May Increase Ecosystem Stability 966
37 Behavioral Biology 893 Inquiry & Analysis 970
Concept 37.1 An Animal’s Genome Influences Its Behavior 894 Retracing the Learning Path 971
Concept 37.2 Learning Also Influences Behavior 896 Assessing the Learning Path 972
Concept 37.3 Thinking Directs the Behavior of Many Animals 897
Concept 37.4 Migratory Behavior Is Both Innate and Learned 898 40 The Living World 974
Concept 37.5 Animal Communication Plays a Key Role in Concept 40.1 Ecosystems Are Shaped by Sun, Wind, and
Ecological and Social Behavior 900 Water 975
Concept 37.6 Natural Selection Shapes Behaviors 903 Concept 40.2 Earth Has 14 Major Terrestrial Ecosystems, Called
Concept 37.7 Behavioral Strategies Have Evolved to Maximize Biomes 979
Reproductive Success 905 Concept 40.3 Freshwater Habitats Occupy Less Than 2% of Earth’s
Concept 37.8 Some Behaviors Decrease Fitness to Benefit Other Surface 981
Individuals 909 Concept 40.4 Marine Habitats Dominate the Earth 982
Concept 37.9 Group Living Has Evolved in Both Insects and Concept 40.5 Humanity’s Pollution and Resource Depletion Are
Vertebrates 912 Severely Impacting the Biosphere 984
Inquiry & Analysis 914 Concept 40.6 Human Activity Is Altering Earth’s Climate 987
38 Ecology of Individuals
and Populations 918 Glossary G1
Concept 38.1 Populations Are Groups of a Single Species in One
Place 919 Appendix A: Answer Key A-1
Concept 38.2 Population Growth Depends upon Members’ Age and Index I-1
Sex 922
xxiv contents
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Title: In the swim

A story of currents and under-currents in gayest New
York
Author: Richard Savage
Release date: September 29, 2023 [eBook #71751]
Language: English
Original publication: Chicago: Rand, McNally & Company, 1898
Credits: Sonya Schermann, David Wilson and the Online

Distributed Proofreading Team at https://www.pgdp.net
(This file was produced from images generously made
available by The Internet Archive)
*** START OF THE PROJECT GUTENBERG EBOOK IN THE SWIM

***
IN THE SWIM.
IN THE SWIM
A Story of
Currents and Under-Currents
in Gayest New York.
By
Richard Henry Savage
Chicago and New York:

Rand, McNally & Company,
Publishers.
Copyright, 1898, by Richard Henry Savage.
All Rights Reserved.
CONTENTS.
BOOK I.
A RISING STAR.
Chapter. Pages
I— “Young
. . Lochinvar
. . . has . .Come
. . out. of. the. West,”
. . . . 5–23
II— The. Drift
. of . a. Day
. in. New
. . York
. City,
. . . . . . . 24–43
III— A Frank
. . Disclosure,
. . . . . . . . . . . . . . 44–67
IV— “Wyman
. . and . . Vreeland”
. . . Swing
. . the. Street,
. . . . . . 68–88
V— Toward
. . the . Zenith,
. . . . . . . . . . . . . 89–110
BOOK II.
WITH THE CURRENT.
VI— In the
. .“Elmleaf”
. . .Bachelor
. . .Apartments,
. . . . . . . . 111–131
VII— “Plunger”
. . .Vreeland’s
. . . Gay
. . Life,
. “Under
. . .the. Rose,”
. . . 132–151
VIII— Miss
. Romaine
. . . Garland,
. . . Stenographer,
. . . . . . . . . 152–170
IX— Senator
. . Alynton’s
. . . Colleague,
. . . . . . . . . . . 171–188
X— An Interview at Lakemere. Some Ingenious
Mechanism.
. . . . “Whose
. . . Picture
. . is . That?”
. . . . . . 189–209
BOOK III.
ON A LEE SHORE.
XI— Miss Marble’s Waterloo! A Lost Lamb! Her Vacant
Chair. Senator Garston’s Disclosure. Sara
Conyers’ Mission. Miss Garland’s Dishonorable
Discharge.
. . . . A .Defiance
. . .to the
. .Death.
. . “Robbed!”
. . . . 210–234
XII— Mine
. and
. .Countermine,
. . . . . . . . . . . . . 235–257
XIII— A Wedding
. . . in. High. .Life,
. . . . . . . . . . 258–279
XIV— For. the. Child’s
. . Sake!
. . . . . . . . . . . . 280–315
XV— In the
. .Dark. Waters,
. . . . . . . . . . . . . 316–361
I N T H E S W I M.
BOOK I—A Rising Star.
CHAPTER I.
“YOUNG LOCHINVAR HAS COME OUT OF THE WEST.”
There was an expression of sullen discontent upon the handsome

features of Mr. Harold Vreeland (gentleman unattached), as the
inbound Hudson River train dashed along under the castled cliffs of
Rhinebeck.
The afternoon was fair—the river of all rivers glittered gaily in the
sun, and a dreamy peace rested on field and stream. But, the peace
of this June afternoon of ’95 entered not into the young wayfarer’s
soul.
The five years which the traveler from nowhere in particular had
thrown away in the far wilds of the sporadic West had not yet robbed
his chiseled features of the good looks which he had borne away
from old Nassau.
And, though his glittering blue eye had been trained to a habitual
impassiveness by much frontier poker, he had always abjured that
Rocky Mountain whisky which “biteth like an adder.”
As he restlessly sought the smoking-car, after a vain struggle with
the all too-evident immorality of a saucy French novel, several
quickly thrilled spinsters followed his retreating form with warm
glances of furtive admiration and half-suppressed sighs.
Vreeland’s stalwart figure was clearly reminiscent of well-played
football and long straining at the oar. His well-set head was bravely
carried, his eye was searching and even audaciously daring in its
social explorations.
At twenty-seven he had not lost the fascination of his soft and
perfectly modulated voice nor the winning insinuation of his too
frequent smile. The chin was far too softly molded for an ascetic, and
an expression of lurking insincerity flickered in the pleasure-loving
curves of his handsome mouth.
But, shapely and glowing with manly vigor, he was a very “proper
man-at-arms” in the battle of life, his sweeping cavalry mustache
lending an air of decision to his sun-burned features.
Though he was perfectly dressed up to the memories of his never-
forgotten “varsity” grade, the “wanderjahre” had given to him a little
of that easy swing which is the gift of wandering on boundless
prairies, long nights spent al fresco under the glittering dome of
stars, and a close commune with the sighing pines of the West.
The shade of bitterness deepened upon his moody face as he noted
a three-masted steam yacht swinging along up the river, with the
elastic quivering throb of her quadruple compounded engines. This
queenly vessel bore the private signal of one American citizen
whose personal finances beggar the resources of many modern
kings.
“Those are the cold-hearted fellows who rule America now with a rod
of iron—the new money kings,” he growled. “Royal by the clink of the
dollar, sovereign by the magic wand of monopoly, impregnable with
the adamantine armor of trusts!”
And then, a lively hatred of the social grandees luxuriously grouped
aft on that splendid yacht crept into his embittered soul.
He could see the Venetian awning which covered the clustered fair-
faced patrician women from the fierce sun, which rudely burns by
day.
And he knew, too, by distant rumors of that superb luxury in which
the American women of the creed of the Golden Calf passed their
happy days in a splendid and serene indolence, only lit up now and
then with gleams of the passion play of high life.
“It’s no use to fight those fellows,” mused Vreeland, as he carefully
trimmed a cigar. “They have come to stay, and I must try and fall into
the train of some one of them.”
He looked back at all those unprofitable years spent beyond the
rugged Rockies. There was a sense of shame and resentment as he
recalled the shabby career of his talented father.
“Thank God, I am now alone in the world, ‘with no one nigh to
hender!’” he bitterly reflected, unconsciously quoting Lowell’s “Zekle
and Huldy.”
The train had rushed on past Poughkeepsie, and the parade music
from West Point floated sweetly across the cool river as the train
halted at Garrison’s for a few moments, before he had morosely
reviewed all the dismal events which brought him a lonely stranger
back to New York.
Erastus Vreeland, a lawyer of no mean accomplishment, had
destined his only son for the bar.
The elder Vreeland was a human spider, who had finally gravitated
downward into the exercise of only the meaner craft of his much-
abused profession.
For long years, in his little office on William Street he had legally
carried on the intrigues of a daring band of clients who rightly should
have ornamented the Academy of Belles-Lettres of New York at Sing
Sing.
During the life of his hoodwinked wife, Vreeland père led a double
existence of more or less moral turpitude, and, at last, a shameless
and successful coup of rascality aroused the ire of a great financial
company.
It was his “notice to quit,” and after the death of his wife, Erastus
Vreeland “swung round the distant circle,” often followed by the
déclassé lawyer.
Omaha, Leadville, Salt Lake, Los Angeles, and other Western cities
finally knew his fox-like cunning and gradually weakening grip.
A political affray, the result of a heated election in Montana, had
been the occasion of the elder Vreeland’s sudden taking off.
And so, the man who had never learned the homely adage that
“corruption wins not more than honesty,” slept far away from his
fathers on the rocky hillsides of Helena, in wild Montana. It was a
miserable summation of failures.
The hegira of the father had left the son stranded in life at the start
upon his finishing the four years at Princeton which had made him
an expert in all the manly arts save any definite plan of money-
getting.
A still self-deceiving man, Erastus Vreeland had hopefully invited his
son to share the suggested exile, and thus, the plan of the law
course for the junior was perforce abandoned. It had not been long
till father and son drifted coldly apart.
The mean, shabby moral nature of the demoralized elder could not
long impose upon the quick-witted youth. The slights of the bench,
the slurs of the bar, the wasp-like thrusts of a bold frontier press, all
proved that the “trail of the serpent” followed on after the talented
weakling whose professional honor was never proof against gold or
gain secured from either side.
And so, with only a hypocritical pretense of a certain lingering
friendly feeling, the two men had finally parted, dividing a few
hundred dollars which were the remains of a retainer in a case,
which deftly went wrong on its trial, sold out, to the benefit of lawyer
Vreeland’s adversary. Then came the bloody finale—and, and—exit
Vreeland pater!
Harold Vreeland sighed in disgust as he recalled the five lost years
of his golden youthful promise.
“It’s all rot,” he muttered, “this idea that the loafer life of the far West
gives either scope, strength, or courage to any man. It is all mere
barbarism, and only a windy discounting of a future which never
comes. A long, bootless struggle with the meaner conditions of life.”
He recalled his varied experiences as notary public, deputy county
clerk, cashier of a shoddy bank—a concern which “folded its Arabian
tents” in six months.
Real estate dealer he had been in several aspiring “boom towns,”
and also, secretary of many frontier “wind” corporations, whose
beautifully engraved stock certificates were now either carried
around in the pocketbooks of dupes or else stuck up in Western
saloons, to the huge edification of the ungodly.
This strange wandering life had made him a fox in cunning, though
not as yet a ravening wolf, for there was little to prey upon in those
dreary distant Occidental preserves. But, his fangs were well
sharpened for the fray.
He realized, as the lights of Haverstraw gleamed out “beyond the
swelling tide,” that he was as yet without any definite plan of
operations.
A singular incident, illustrative of the roughly good-humored social
code of the wild West had caused him to seek the city of Manhattan.
The political clique which had coolly plotted the murder of his crafty
father, with a last generous twinge of conscience, had sent all the
private papers of the defunct lawyer over to his son, who was
listlessly engaged at the time in endeavoring, on a net cash capital of
a hundred and fifty dollars, to float a ten million dollar corporation, in
order to utilize certain waste energy of those foaming falls of the
Spokane River, which have so long caused both the salmon and the
Indians a great deal of unnecessary trouble.
And then, young Vreeland wearily explored those ashes of life—the
“papers in the case” of the defunct.
The unwelcome discovery of many evidences of his father’s shame
and the revealing of all that secret life which had sent his patient
mother to the shadowy bourne of heartbroken wives, was somewhat
mitigated by the discovery of a paid-up policy of ten thousand dollars
in the great “Acqueduct Life Insurance of New York City.”
There was, as usual, some strings and filaments hanging out loosely
knotted up, and it had been a labor of months, involving a
correspondence of some acerbity, for him to obtain letters of
administration, close up his father’s “estate,” and depart to Gotham
to receive a check for seven thousand dollars in full settlement of the
claim.
On the road over from Spokane, Mr. Harold Vreeland had carefully
counted all his ships. He had even gone over all his own abortive
attempts at opening any useful career, and so, on this summer
evening, he gloomily felt how poorly prepared he was to fight the
battle of life against the keen competition and increasing pressure of
his peers in New York City.
“If I had only my father’s profession, I would have a chance to get in
among these fellows, and I would soon have my share of the gate
money,” he growled.
“But to take a place in the line of mere drudges, to sink down into the
death in life of a hall room and a cheap boarding-house. Once
planted there, I am there forever. And I have not a friend in the whole
world!”
His mental harvest had only been one of husks, and he keenly felt
the absence of any definite calling pour accrocher.
Suddenly his eye caught the gleam of a sunset upon a dozen
drifting, glittering white sails on the river.
They all seemed to float on serenely, borne along upon the broad
tide, with no visible man’s hand to guide.
“I will drift a while,” he murmured. “I have a few thousand dollars.
Something will surely turn up. If it does not,” he resolutely said,
“then, I will turn it up myself.”
“There are women here, too—women with hearts of flame, and who
are to be won. I was a fool ever to go out to the frontier. Perhaps—”
And his mind reverted to a lucky college chum who had married a
woman nearly two generations older than himself, but a well-
preserved Madame “Midas.”
“By Jove! anything is better than this beastly poverty,” he mused.
“Even that.”
“This is no era for poor men. Poverty is the only crime nowadays.”
His cynicism was broken off by the approach of two men, who rose
to rejoin friends in the train as it dashed along toward the Bronx
River.
As they came up the smoking-car, Vreeland easily recognized Fred
Hathorn, the stroke of the college crew in which he had once won
hard-fought honors for the orange.
There was no mistaking the easy luxury which exhaled from Mr. Fred
Hathorn of the great firm of Hathorn and Potter, bankers and brokers
of dingy Wall Street, a man who had already arrived!
The first crucial glance of rapid inspection was not lost on Vreeland,
as Hathorn, in an easy way cried: “Hello, Hod Vreeland! What brings
you over here?”
With a perfunctory politeness, Mr. James Potter halted and calmly
acknowledged Hathorn’s listless introduction.
The little blonde man-about-town, however, gazed longingly ahead at
the car where certain fair dames now awaited their escorts.
Jimmy Potter was born to “no end of easy money,” and so his
dashing senior partner’s genius for finance was strongly buttressed
by the whirlwind of cash which clustered around Jimmy Potter’s
lucky head.
All sorts of financial bees seemed to swarm around Potter and
quietly settle in his hive.
“What’s the use of making a row?” he often remarked. “Sit still, and
what you want in life will come to you.” Mr. James Potter of New York
was an Epicurean disciple.
The blood mounted to Vreeland’s forehead as he noted all the
deprecating courtesy of Hathorn’s welcome.
“Damn him! I’ll give him a bit of a bluff,” he quickly decided, under
the inspiration of some bold, familiar spirit.
There was an air of quiet comfort in the careless response of
Vreeland.
“I have just fallen into a good bit of money by my father’s death, and
so have come on here to enjoy myself. I may spend a couple of
years abroad.”
Vreeland then blessed that daring, familiar spirit which so saucily
suggested his “cheeky” retort, as the man who had been his chum
and fellow of several Greek letter societies stopped short. “Wait for
me at the station, old fellow. We are bothered yet with some ladies.
They leave at the station. Then we will dine later at the club and talk
over old times a bit. You’ll come, too, won’t you, Potter?”
Jimmy Potter carelessly nodded an assent from sheer laziness, and
then the two members of the jeunesse dorée, passed on into the
boudoir car.
There was a twinkle of triumph in Vreeland’s eye as he sank back in
his seat.
“I got a dinner out of you at any rate, Mr. Snob,” he gleefully
chuckled.
And, highly elated, he decided then and there, to vary his first plan of
drifting with the tide, and to cautiously put his oar in a bit where it
would help him on.
His step was as light as the tread of a panther when he leaped out of
the car at Forty-second Street.
“I’ll have a stolen glance at their women,” he quickly resolved.
“Perhaps they may give dinners, too.”
And just then, there seemed to be the twinkle of a little star of Hope
lighting up that devious, unknown path which he was so soon to
tread.
“I’ll let him give me a Club card,” he mused, as the wearied
passengers hurried along to brave the din of importunate jehus.
He was wondering how much of a social show he could make at
need with his slender fortune, when the two men slowly approached
with three “shining ones” of the golden strata of womanly New York.
“These people are all in the swim,” he murmured. “I will find the way!
I am as good as any of them.”
And as he raised his eyes, he met the glances of the imperial-looking
woman who was Fred Hathorn’s companion.
The lady’s eyes rested for a moment upon the handsome stranger,
and then fell with a peculiar abruptness.
“If that woman plays any star part in his life, I will try and take her
away from him,” resolved Vreeland, whose whole soul was now
thrilling with the beautiful woman’s sudden, startled admission of
interest in a passing stranger. The wine of life stirred in the young
wanderer’s veins.
His audacious, familiar sprite suggested the profound bow which
was Mr. Harold Vreeland’s first salutation upon the outskirts of the
“Four Hundred.”
He had adroitly managed to convey the respectful homage of the
salutation by his velvety eyes to the very person intended, for, while
Jimmy Potter was placidly listening to the brilliant chatter of two very
vivacious rosebuds, Mrs. Elaine Willoughby turned to Hathorn:
“Fred, who is your Western friend?” she asked, with an assumed
carelessness.
It was by sheer good luck that Hathorn, who was watching the young
millionairess whom he was soon to marry, answered with an unusual
warmth:
“An old college chum—Vreeland of Princeton, and a rattling good
fellow.”
Fred Hathorn eyed with a certain qualminess the easy aplomb of his
Crœsus partner, as Jimmy Potter pressed closely to the side of
Hathorn’s destined bride, Miss Moneybags.
That young lady was destined to play the rôle of Queen of Diamonds
in the ambitious young banker’s life.
He had resolutely set up the motto, “Aut Cæsar, aut nullus,” and he
was just a bit shy of the beloved James trifling with his dashing
fiancée.
“All sorts of things happen in New York,” mused the agnostic
Hathorn, as he handed the ladies into a waiting victoria and then
turned to rejoin the man who more than ever had now decided to
paddle a bit, as well as to drift on with the tide of fortune.
There was a glow of satisfaction burning in the Western adventurer’s
heart as, half an hour later, he noted Hathorn dash off his potent
signature behind his guest’s name on the visitor’s book of the Old
York Club. It was the open sesame to the regions of the blest—
young New York par excellence.
The trio adjourned to the billiard room, and, then and there, Vreeland
for the first time tasted the famous club cocktail.
He was “living up to his blue china,” as he gravely bowed when
Hathorn gave him a two-weeks’ card.
“I’ll have it renewed for you, old fellow,” lightly remarked the young
banker.

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changes to this edition vii

The Learning Path

Bryophytes Have a Dominant

Every chapter opensAutorhythmic Cells Contraction is initiated by the SA

main concepts in the Angiosperms Are

Assessing the Learning Path

Evolution Explains the Unity and

Ferns and Allies

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Uncovering Connections Between Concepts

• Early life did not use oxygen.

• Life descended from early cells that spontaneously

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make ATP and NADPH Living systems Living systems

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• Photosystems are erm

important structures that

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xvi a learning path to understanding biolog y

PART I The Molecular Concept 4.3

14 DNA: The Genetic Material 280 17 Biotechnology 358

23 Prokaryotes and Viruses 497

Title: In the swim

Author: Richard Savage

Release date: September 29, 2023 [eBook #71751]

Original publication: Chicago: Rand, McNally & Company, 1898

Credits: Sonya Schermann, David Wilson and the Online