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Biology
the dynamic science
seco
se
e co
c o nd
n eedi
d i ti
t on
n
RUSSELL HERTZ McMILLAN
Volume 1
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1 2 3 4 5 6 7 14 13 12 11 10
Peter J. Russell received a B.Sc. in Biology from the University of Sussex, England, in 1968 and a
Ph.D. in Genetics from Cornell University in 1972. He has been a member of the Biology faculty of
Reed College since 1972; he is currently a professor of biology. Peter teaches a section of the intro-
ductory biology course, a genetics course, and a research literature course on molecular virology. In
1987 he received the Burlington Northern Faculty Achievement Award from Reed College in recog-
nition of his excellence in teaching. Since 1986, he has been the author of a successful genetics text-
book; current editions are iGenetics: A Molecular Approach, iGenetics: A Mendelian Approach, and
Essential iGenetics. Peter’s research is in the area of molecular genetics, with a specific interest in
characterizing the role of host genes in the replication of the RNA genome of a pathogenic plant
virus, and the expression of the genes of the virus; yeast is used as the model host. His research has
been funded by agencies including the National Institutes of Health, the National Science Founda-
tion, and the American Cancer Society. He has published his research results in a variety of journals,
including Genetics, Journal of Bacteriology, Molecular and General Genetics, Nucleic Acids Research,
Plasmid, and Molecular and Cellular Biology. Peter has a long history of encouraging faculty research
involving undergraduates, including cofounding the biology division of the Council on Undergradu-
ate Research in 1985. He was Principal Investigator/Program Director of a National Science Founda-
tion (NSF) Award for the Integration of Research and Education to Reed College, 1998–2002.
Paul E. Hertz was born and raised in New York City. He received a B.S. in Biology from Stanford
University in 1972, an A.M. in Biology from Harvard University in 1973, and a Ph.D. in Biology from
Harvard University in 1977. While completing field research for the doctorate, he served on the Biology
faculty of the University of Puerto Rico at Rio Piedras. After spending two years as an Isaac Walton
Killam Postdoctoral Fellow at Dalhousie University, Paul accepted a teaching position at Barnard
College, where he has taught since 1979. He was named Ann Whitney Olin Professor of Biology in
2000, and he received The Barnard Award for Excellence in Teaching in 2007. In addition to serving on
numerous college committees, Paul chaired Barnard’s Biology Department for eight years. He is also
the Program Director of the Hughes Science Pipeline Project at Barnard, an undergraduate curriculum
and research program that has been funded continuously by the Howard Hughes Medical Institute
since 1992. The Pipeline Project includes the Intercollegiate Partnership, a program for local commu-
nity college students that facilitates their transfer to four-year colleges and universities. He teaches one
Aaron W. Kinard
semester of the introductory sequence for Biology majors and preprofessional students, lecture and
laboratory courses in vertebrate zoology and ecology, and a year-long seminar that introduces first-year
students to scientific research. Paul is an animal physiological ecologist with a specific research interest
in the thermal biology of lizards. He has conducted fieldwork in the West Indies since the mid-1970s,
most recently focusing on the lizards of Cuba. His work has been funded by the NSF, and he has pub-
lished his research in such prestigious journals as The American Naturalist, Ecology, Nature, Oecologia,
and Proceedings of the Royal Society. In 2010, he received funding from NSF for a project designed to
detect the effects of global climate warming on the biology of Anolis lizards in Puerto Rico.
Beverly McMillan has been a science writer for more than 25 years and is coauthor of a college
text in human biology, now in its eighth edition. She has worked extensively in educational and
commercial publishing, including eight years in editorial management positions in the college divi-
sions of Random House and McGraw-Hill. In a multifaceted freelance career, Bev also has written
or coauthored 10 trade books, as well as story panels for exhibitions at the Science Museum of
Virginia and the San Francisco Exploratorium. She has worked as a radio producer and speech-
writer for the University of California system and as a science writer and media relations advisor
for the Virginia Institute of Marine Science of the College of William and Mary. She holds under-
graduate and graduate degrees from the University of California, Berkeley.
vi
vii
viii P R E FA C E
P R E FA C E ix
8
study outline ❮ Study Outline The Study Outline
provides an overview of all the topics
8.1 Overview of Cellular
and key concepts in the chapter. Each
155
❯ Study Break Encourages you to pause and think STUDY BREAK 8.2 <
about the key concepts you have just encountered 1. What are the energy-requiring and energy-releasing steps of
glycolysis?
before moving to the next section. 2. What is the redox reaction in glycolysis?
3. How is ATP synthesized in glycolysis?
4. Why is phosphofructokinase a target for inhibition by ATP?
xiii
Hot Potatoes: Do Plants Use 3. Laloi and her coworkers then tested stressed by low temperatures. Thus, mecha-
exciting and interesting research relying on molecular techniques;
Uncoupling Proteins to whether exposing potato plants to cold nisms for warming body tissues once thought
Generate Heat?
Mammals use several biochemical and
temperatures could induce greater synthe-
sis of the UCP mRNA (Figure 2). The cold
treatment resulted in an increase in UCP
to be the province only of animals appear to
be much more widespread. In particular,
UCPs, which were believed to have evolved in
these molecular tools allow researchers to answer questions that
molecular processes to maintain body heat.
One process is shivering; the muscular
activity of shivering releases heat that helps
mRNA level in the leaves.
Conclusion
The research indicates that potato plants
relatively recent evolutionary times with the
appearance of birds and mammals, may be a
much more ancient development.
they could not even pose 20 or 30 years ago.
keep body temperature at normal levels.
Another mechanism operates through un- probably use the mitochondrial uncoupling Source: M. Laloi et al. 1997. A plant cold-induced
coupling proteins (UCPs), which eliminate process to warm tissues when they are uncoupling protein. Nature 389:135–136.
the mitochondrial H⫹ gradient by making the
inner mitochondrial membrane leaky to
Flowers—very high
protons. Electron transfer and the oxidative
UCP mRNA level
reactions then run at high rates in mitochon-
❯ Focus on Research Special boxes in most chapters present research topics in more depth.
spots
p are identified byy comparing
p g their locations with identifyy the molecules that
are exposed to a reactant chemical that has been
the radioactivity to produce an imag image of the
the positions of spots made by known molecules when were radioactive. “labeled” with a radioactive isotope such as 14C
gland on a photo
photographic
graphic film Examples
lm. Exa m of what
the “knowns” are run through the same procedure. or 3H. After being exposed to the tracer, the
presented
the scans may show are presente nted in the figure.
chemical products in which the isotope appears,
Another application uses the factt tha that radioactive
and their sequence of appearance, can be de-
regions
thallium is not taken up by region ns oof the heart
FOCUS ON Model Research Organisms tected by their radioactivity and identifi fied.
identified.
muscle with poor circulation to dete detect coronary
For example, algae and plants use carbon di-
artery disease. Other isotopes aree us used to detect
Normal Enlarged Cancerous
The Yeast Saccharomyces oxide (CO2cul-
The cells growing in Saccharomyces ) as a raw material
mammals.in photosynthesis.
Genetic studies with Saccharomyces
bone injuries and defects, including injured, ar-
To trace Ifthe reactions ofwere
photosynthesis,
also the firstMelvin Scans of human thyroid glands after iodine-123
cerevisiae tures in the research lab are haploid. the to show the thritic,
genes orcarried in
abnormally growing segsegments
gme of bone.
was injected into the bloodstream. The radioactive
culture conditions are kept at Calvin
optimaland his coworkers
levels thegrew
DNAalgal cells in a
of mitochondria and their patternsof
Treatment ofdisease with radiois
radioisotopes takes
Saccharomyces cerevisiae, commonly known as iodine becomes concentrated in the thyroid gland.
(which requires only a source medium with CO2 that inheritance.
of a fermentable contained the radioiso-
The complete DNA sequenceofof
advantage radioactivity
the fact that radioac ctivi in large
baker’s yeast or brewer’s yeast, was probably
sugar such as glucose, a nitrogen source, and S. cerevisiae, which includes more than 12 mil-
the first microorganism to have been grown
minerals), the cells reproduce asexually by lion base pairs that encode about 6,000 genes,
and kept in cultures—a beer-brewing vessel is
budding. Saccharomyces has two mating was the first eukaryotic genome to be obtained.
basically a Saccharomyces culture. Favorite
types. If two yeast cells of different mating Analysis of the genome sequence revealed that
strains of baker’s and brewer’s yeast have been
types contact one another, they fuse—mate— yeast has many genes related to those in ani-
kept in continuous cultures for centuries. The
producing a diploid cell. Diploid cells can also mals, including mammals, making this rela-
yeast has also been widely used in scientific re-
reproduce asexually by budding. Under cer- tively simple microorganism an excellent sub-
search; its microscopic size and relatively
tain conditions, diploid yeast cells undergo ject for research that can be applied to the
short generation time make it easy and inex-
meiosis, producing haploid spores which are more complex animals of interest.
pensive to culture in large numbers in the
laboratory.
resistant to harsh conditions in the environ-
ment. Yeast spores germinate into haploid
cells, which then reproduce asexually.
The genes of yeast can also be manipu-
lated easily using genetic engineering tech-
niques. This has made it possible for re-
Focus on Model Research Organisms explains
Genetic studies with Saccharomyces led to
the discovery of some of the genes that control
searchers to alter essentially any of the yeast
genes experimentally to test their functions why researchers use certain organisms as
Adrian Warren/Ardea/London
the eukaryotic cell cycle. Many of these genes, and to introduce genes or DNA samples from
after their first discovery in yeast cells, were other organisms for testing or cloning. Sac-
found to have counterparts in animals and
plants. Defective versions of the genes often
charomyces has been so important to genetic
studies in eukaryotes that it is often called the
research subjects.
contribute to the development of cancer in eukaryotic E. coli.
xiv
Protocol:
Observations
Deductive
FIGURE 1.16 Observational Research reasoning
3. Use deductive reasoning to make predictions about what you would observe if
the hypothesis were applied to a novel situation.
A Field Study Using a Null Hypothesis Hypothesis: Anolis cristatellus, the crested anole, uses patches of sun and shade to regulate
its body temperature. Predictions
Null Hypothesis: Because this lizard does not regulate its body temperature, individuals se-
Observational Research lect perching sites at random with respect to environmental factors that influence body
temperature.
Anolis cristatellus Method: The researchers created a set of hollow, copper lizard models, each equipped with 4. Design and conduct a controlled experiment (or new observational study) to
a temperature-sensing wire. At study sites where the lizards live in Puerto Rico, the research- test the predictions of the hypothesis. The experiment must be clearly defined
ers hung 60 models at random positions in trees. They observed how often live lizards and so that it can be repeated in future studies. It must also lead to the collection of
the randomly positioned copper models were perched in patches of sun or shade, and they Experiments measurable data that other researchers can evaluate and reproduce if they
measured the temperatures of live lizards and the copper models. Data from the randomly choose to repeat the experiment themselves.
positioned copper models define the predictions of the null hypothesis.
Results: The researchers compared the frequency with which live lizards and the copper
models perched in sun or shade as well as the temperatures of live lizards andd thee copper
c
models. The data revealed that the behavior and temperatures of A. cristatelluss we
ere differ-
were IInterpreting
Interpret
t t ing
i the
th l t Compare the results of the experiment or new observations with those predicted by the hypothesis. Scientists
h Results:
R esults: Sc often use
origginal
ent from those of the randomly positioned models, therefore confirming the original formal statistical tests to determine whether the results match the predictions of the hypothesis.
FIGURE 8.15 Experimental Research
Alejandro Sanchez
hypothesis.
The Racker and Stoeckenius Experiment Question: Does an H⫹ gradient power ATP synthase-catalyzed ATP synthesis, thereby sup-
Copper Anolis model Percentage of models and Temperatures of ATP Synthesis by ATP Synthase Experiment: Efraim Racker of Cornell University and Walther Stoeckenius of University of
lizards perched in sun or shade models and lizards California, San Francisco, made membrane vesicles that had a proton pump and ATP syn-
Kevin de Queiroz, National Museum of Natural History, Smithsonian
Anolis cristatellus Anolis cristatellus thase to determine whether proton-motive force drives ATP synthesis.
100 30 1. The researchers constructed synthetic
Percentage in sun or shade
Percentage of observations
Perched in shade Temperature ( oC) illuminated the vesicles and then ana-
lyzed the concentration of H⫹ in them.
Conclusion: A. cristatellus uses patches of sun and shade to regulate its body temperature. Cutaway view of a synthetic vesicle
Source: P.E. Hertz. 1992. Temperature regulation in Puerto Rican Anolis lizards: A field test using null hypotheses. Ecology 73:1405–1417.
Result: H⫹ is pumped into the vesicles, creating an H⫹ gradient.
ADP
ATP ADP
synthase Headpiece end
ADP + P ATP
i
Result: In the dark, no ATP was synthesized. Result: In the light, ATP was synthesized.
Together, these results showed that light activated the bacteriorhodopsin to produce an
H⫹ gradient, with H⫹ moving from the outside to the inside of the vesicle (like the move-
ment from the mitochondrial matrix to the intermembrane compartment in Figure 8.13),
and that the energy from the H⫹ gradient drove ATP synthesis by ATP synthase.
(5-FU) and cisplatin (CDDP) are drugs that work in this way. 5-FU 60 Schedule B: CDDP (8h)
help you build your skills in analyzing figures and in inhibits DNA replication, while CDDP binds to DNA causing changes
that cannot be corrected by DNA repair enzymes so that programmed
followed by 5-FU
Schedule C: 5-FU and CDDP
cell death (see Chapter 7) is triggered. Researchers suspected that 40
reading graphs and tables. these drugs might be useful in treating human gastric cancer and
(8h) followed by 5-FU
Schedule D: 5-FU (8h), 5-FU
tested their effectiveness in gastric cancer cells growing in culture. and CDDP (8h), followed
20
They added 5-FU alone, or 5-FU and CDDP in various timed combi- by 5-FU
nations (schedules) to cultured gastric cancer cells and measured the Schedule E: 5-FU (16h), 5-FU
inhibitory effects of the drugs on cell proliferation compared with and CDDP (8h), followed
untreated cells. The results in the figure to the right shows for each A B C D E
by 5-FU
schedule the % cell proliferation, meaning the proliferation of treated Schedule
cells/proliferation of control, untreated cells ⫻ 100%.
1. Which drug schedule was the most effective? Source: H. Cho, et al. 2002. “In-vitro effect of a combination of
2. How did the drug schedule in the most effective treatment differ 5-fluorouracil (5-FU) and cisplatin (CDDP) on human gastric cancer
from the schedules in all the other treatments? cell lines: timing of cisplatin treatment.” Gastric Cancer 5:43–46.
>
❯ Think Outside the Book These individual, THINK OUTSIDE THE BOOK
team, or Internet research activities encourage active Scientists have been working to develop an arti-
ficial version of photosynthesis that can be used
learning and peer discussion. They foster critical to produce liquid fuels from CO2 and H2O. Col-
laboratively or individually, find an example of
thinking skills by engaging you in the topic. research on artificial photosynthesis and pre-
pare an outline of how the system works or is
anticipated to work.
xv
the major points. Orientation diagrams are inset on figures and Pyruvate
Inner
mitochondrial
two electron carriers, FMN (flavin
mononucleotide) and an Fe/S
(iron–sulfur) protein, to
chromes a and a3 to oxygen. Four
protons are added to a molecule
of O2 as it accepts four electrons,
7 Because of the gradient, H+
oxidation
flows across the inner membrane
help you identify the specific biological process being depicted membrane ubiquinone. forming 2 H2O. and into the matrix through a
channel in the ATP synthase.
Citric
2 Complex II oxidizes FADH2 to 5 As electrons move through
and where the process takes place. acid
cycle
ATP FAD; the two electrons released
are transferred to ubiquinone, and
the electron transfer system, they
release free energy. Part of the
8 The flow of H+ activates ATP
synthase, making the headpiece
the two protons released go into released energy is lost as heat, and stalk rotate.
the matrix. Electrons that pass to but some is used by the mito-
ubiquinone by the complex II chondrion to transport H+ across
Oxidative 9 As a result of changes in
ATP reaction bypass complex I of the the inner mitochondrial
phosphorylation shape and position as it turns, the
electron transfer system. membrane from the matrix to the
headpiece catalyzes the synthesis
inter membrane compartment at
of ATP from ADP and Pi.
3 Complex III accepts electrons complexes I, III, and IV.
from ubiquinone and transfers
them through the electron carriers
in the complex—cytochrome b, an
Image copyright javarman, 2010. Used under
ments of chloroplasts.
license from Shutterstock.com
Outer
mitochondrial
membrane
6
H+ H+ High H+
H+ H+
H+ H+ H+ H+ H+
H+ H+ H+
Cutaway of a small One of the photosynthetic H+ H+ H+ H+ H+
H+
section from the leaf cells, with green chloroplasts H+
ATP
synthase Stator
Leaf’s upper surface Photosynthetic cells Intermembrane
compartment
cyt c
Large central cyt b e– 4
4 Basal
Basal
vacuole e– 3 3
3 cyt a e– cyt a3 7 unit
unit
e– e–
Inner FMN e– 1 1 Ubiquinone Fe/S
3 8
mitochondrial Fe/S e– e– e–
(CoQ) Complex 4
membrane e– Complex
cyt c1 IV Stalk
Complex e– III
5 H2O
I
1 Complex 2 H+
II
5 H+
CO2 5
NADH + H+ H+ 2 H+ + 1/ O2
H+
2
Head-
piece
NAD+ FADH2 FAD + 2 H+
Mitochondrial Low H+
matrix 6
O2 9
Stomata (through Nucleus
ADP + P ATP
i
which O2 and CO2
are exchanged with Electron transfer system Chemiosmosis
the atmosphere) Electrons flow through a series of proton (H+) ATP synthase catalyzes ATP
pumps; the energy released builds an H+ gradient synthesis using energy from the H+
across the inner mitochondrial membrane. gradient across the membrane.
Oxidative phosphorylation
Cutaway view of a chloroplast
FIGURE 8.13
Oxidative phosphorylation: The mitochondrial electron transfer system and chemiosmosis. Oxidative phosphorylation involves the electron transfer
Outer membrane 1system
66 (stepsU N1–6),
I T Oand
N E chemiosmosis
M O L E C U L by
E SATP
A Nsynthase
D C E L L(steps
S 7–9). Blue arrows indicate electron flow; red arrows indicate H⫹ movement.
Inner membrane
Granum
xvi
Test Your Knowledge 4. Which of the following statements is false? Imagine that you in-
1. What is the final acceptor for electrons in cellular respiration? gested three chocolate bars just before sitting down to study this
a. oxygen chapter. Most likely:
b. ATP a. your brain cells are using ATP.
c. carbon dioxide b. there is no deficit of the initial substrate to begin glycolysis.
d. hydrogen c. the respiratory processes in your brain cells are moving
e. water atoms from glycolysis through the citric acid cycle to the
electron transfer system.
2. In glycolysis: d. after a couple of hours, you change position and stretch to ❮ Understand and Apply End-of-chapter
a. free oxygen is required for the reactions to occur. rest certain muscle cells, which removes lactate from these
b. ATP is used when glucose and fructose-6-phosphate are muscles. questions focus on factual content in the
phosphorylated, and ATP is synthesized when 3-phospho- e. after 2 hours, your brain cells are oxygen-deficient.
glycerate and pyruvate are formed.
5. If ADP is produced in excess in cellular respiration, this excess chapter while encouraging you to apply
c. the enzymes that move phosphate groups on and off the
ADP will:
molecules are uncoupling proteins.
a. bind glucose to turn off glycolysis. what you have learned.
d. the product with the highest potential energy in the pathway
is pyruvate. b. bind glucose-6-phosphate to turn off glycolysis.
e. the end product of glycolysis moves to the electron transfer c. bind phosphofructokinase to turn on or keep glycolysis
system. turned on.
d. cause lactate to form.
3. Which of the following statements about phosphofructokinase is e. increase oxaloacetate binding to increase NAD+ production.
false?
a. It is located and has its main activity in the inner mitochon- 6. Which of the following statements is false? In cellular respiration:
drial membrane. a. one molecule of glucose can produce about 32 ATP.
b. It catalyzes a reaction to form a product with the highest b. oxygen combines directly with glucose to form carbon dioxide.
potential energy in the pathway. c. a series of energy-requiring reactions is coupled to a series of
c. It can be inactivated by ATP at an inhibitory site on its energy-releasing reactions.
surface. d. NADH and FADH2 allow H+ to be pumped across the inner
d. It can be activated by ADP at an excitatory site on its surface. mitochondrial membrane.
e. It can cause ADP to form. e. the electron transfer system occurs in the inner mitochon-
drial membrane.
Express Your Opinion Express Your Opinion encourages you to weigh both sides
Developing new drugs is costly. There is little incentive for pharma- of an issue by reading pro/con articles, and then make your
ceutical companies to target ailments that affect relatively few indi-
viduals, such as Luft syndrome. Should governments allocate some opinion known through an online voting process.
funds to private companies that search for cures for rare disorders? Go
to CengageNOW to investigate both sides of the issue and then vote
online.
xvii
xviii
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