Cellular Respiration

Cellular Respiration

A. Malcolm Campbell, PhD

Christopher J. Paradise, PhD
Cellular Respiration
Copyright A. Malcolm Campbell and Christopher J. Paradise. 2016.

All rights reserved. No part of this publication may be reproduced, stored

in a retrieval system, or transmitted in any form or by any means
electronic, mechanical, photocopy, recording, or any other except for
brief quotations, not to exceed 250 words, without the prior permission
of the publisher.

First published in 2016 by

Momentum Press, LLC
222 East 46th Street, New York, NY 10017
www.momentumpress.net

ISBN-13: 978-1-60650-997-5 (print)

ISBN-13: 978-1-60650-998-2 (e-book)

Momentum Press Biology Collection

Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

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10 9 8 7 6 5 4 3 2 1

Printed in the United States of America

 Abstract
What happens to a meal after it is eaten? Food consists primarily of
lipids, proteins and carbohydrates (sugars). How do cells in the body
process food once it is eaten and turned it into a form of energy that
other cells can use? This book examines some of the classic experimen-
tal data that revealed how cells break down food to extract the energy.
Metabolism of food is regulated so that energy extraction increases when
needed and slows down when not needed. This type of self-regulation
is all part of the complex web of enzymes that convert food into energy.
Adding to this complexity is that all food eventually winds up as two
carbon bits that are all processed the same way. This book will also reveal
why animals breathe oxygen and how that relates to the end of the en-
ergy extraction process and oxygens only role in the body. Rather than
look at all the details, this book takes a wider view and shows how cel-
lular respiration is self-regulating.

Keywords
first law of thermodynamics, second law of thermodynamics, entropy,
free energy, reduced, oxidized, homeostasis, palmitic acid, casein, lactose,
enymes, redox, cofactors, coenzymes, beta oxidation, acetyl CoA, mitro-
chondrial matrix, cellular respiration, limiting factor, deamination, alloste-
rically modulated, phosphofructokinase, citrate, negative feedback loop,
mitochondrial matrix, chemiosmosis, ATP synthase
 Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Molecules Carry Energy in their Covalent Bonds...............1
Chapter 2 Converting Common Foods into Energy.........................11
Lipid Metabolism.............................................................13
Protein Metabolism..........................................................20
Carbohydrate Metabolism................................................25
Ethical, Legal, Social Implications:
Memorizing Details Obscures Learning........................32
Chapter 3 Energy Extraction from 2-Carbon Intermediates..............39
Chapter 4 ATP Production from Digested Foods.............................47
Overview of Energy Homeostasis.....................................54
Ethical, Legal, Social Implications:
The Importance of Eating Vegetables............................57
Conclusion............................................................................................61
Glossary................................................................................................63
Index....................................................................................................65
 Preface
This book on cellular respiration is part of a thirty book series that col-
lectively surveys all of the major themes in biology. Rather than just pres-
ent information as a collection of facts, the reader is treated more like a
scientist, which means the data behind the major themes are presented.
Reading any of the thirty books by Campbell and Paradise provides read-
ers with biological context and comprehensive perspective so that readers
can learn important information from a single book with the potential to
see how the major themes span all size scales: molecular, cellular, organ-
ismal, population and ecologic systems. The major themes of biology en-
capsulate the entire discipline: information, evolution, cells, homeostasis
and emergent properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
Readers will learn, in an engaging and very non-conventional way, how
cells extract energy from lipids, proteins and carbohydrates and some
of the supporting evidence behind our understanding. The historic and
more recent experiments and data will be explored. Instead of believing
or simply accepting information, readers of this book will learn about the
science behind cellular respiration the same way professional scientists
dowith experimentation and data analysis. In short, data are put back
into the teaching of biological sciences.
Readers of this book who wish to see the textbook version of this content
can go to www.bio.davidson.edu/icb where they will find pedagogically-
designed and interactive Integrating Concepts in Biology for introductory
biology college courses or a high school AP Biology course.
 Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with AMC. Davids gift allowed us to hire talented artists (Tom Webster
and his staff at Lineworks, Inc.) and copyeditor Laura Loveall. Thanks go
to Kristen Mandava for project management and guidance on the pub-
lishing process. In particular, we are indebted to Katie Noble and Melissa
Hayban for their many hours of help and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to admin-
istrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond, Verna
Case, and Barbara Lom who had confidence in us and encouraged us to
persist despite setbacks along the way.
These books were the product of the shared labor of my two vision-
ary coauthors Laurie Heyer and Chris Paradise. We shared the dream and
the hardships and developed this book from scratch. My family has been
very supportive and I thank Susan, Celeste and Paulina for their support
and patience. I also want to thank Jan Serie, my pedagogical mentor, who
taught me so much about the art and science of helping students learn.
I benefited from the support of the Howard Hughes Medical Institute
grant 52006292, the James G. Martin Genomics Program, and Davidson
College. This book would not have survived its first draft without my
students who endured the typos and the early versions of this book. These
undergraduates participated in a bold experiment to see if beginners could
construct their own knowledge, retain what they learned, and transform
the way they see themselves and the discipline of biology. While many
people said that beginning students were not up to the task, my students
proved them wrong.
 Introduction
Living takes energy and energy comes from the food we eat. How is a
sandwich converted into the molecules that muscles use to chew the
sandwich? It is hard to comprehend many molecular processes, especially
energy extraction. Energy cannot be seen, tasted or smelled and yet its
properties are experienced all the time. This book reveal how molecules
in a persons cells convert the eaten food into energy that can be used by
cells. This exploration begins with a review of some chemistry terms. It
is important to remember how molecular energy is stored in chemical
bonds. It is also important to remember the basics of how energy is mea-
sured and described by chemists. Chemistry is a foundational science that
helps make biology understandable, especially at the molecular level. This
book focus on how cellular metabolism gradually breaks bigger molecules
into smaller ones while transferring the energy from the bonds in food
to bonds in molecules used to transport energy between and within cells.
Ultimately, cells produce molecules of ATP which can be used for many
functions. Along the way, cellular metabolism produces waste products
containing carbon and oxygen atoms as well as wasted energy cells were
unable to harness.
 CHAPTER 1

Molecules Carry Energy

in their Covalent Bonds

Everyone has watched water boil, and perhaps over an open fire. From
personal experience it is clear that burning wood releases a lot of heat
energy, but the molecular source of this energy may not be clear, nor is
how the energy is released. Notice that with a campfire, some of the heat
is radiated to the air around the fire, and the remaining heat is absorbed
by the metal pot and eventually the water to produce steam. Convert-
ing liquid water into gaseous steam is work, as defined in the chemical
sense. Converting matter from one state into another is a type of work
that requires an input of energy, such as heat from the fire. It is possible
to harness steam to do other work, which is what happens inside coal
and nuclear power plants. Steam spins paddles of a turbine, which causes
a magnet inside coiled wires to spin. A spinning magnet inside coiled
wires produces a flow of electrons (e-), which is electricity. Wind genera-
tors and hydroelectric dams also produce a flow of electrons by spinning
magnets inside wire coils. Solar panels produce electron flow from un-
stable compounds that absorb the suns energy. In short, electrons are a
fundamental form of energy that are consumed every time an electrical
device is turned on.
There are two physical laws related to energy that apply to everything
in the universe. The field of study that focuses on the conversion of heat
to mechanical work is called thermodynamics, coming from the Greek
terms for heat (thermo) and power (dynamics). The field of thermody-
namics has two laws that are pertinent to understanding of molecular
energy sources used to do work. The first law of thermodynamics states
that energy can be transformed (changed from one form to another) but
2 CELLULAR RESPIRATION

cannot be created or destroyed. The second law of thermodynamics

states that the entropy, or disorder, of a system increases over time unless
energy is expended to reduce entropy. Biology has many hypotheses, a
few of which have been supported often enough that they are elevated to
the status of theories. In physics, theories that have been demonstrated to
be fundamental to all actions within the universe are called laws to signify
their fundamental roles in nature.
In addition to the boiling water, campfires produce smoke that con-
tains CO2, water vapor, ashes, and lost heat, all of which are byproducts
of releasing the energy from the wood. The match used to start the fire
supplied enough energy to break a few covalent bonds of the wood and
produce CO2 and H2O from oxygen and the wood. The new covalent
bonds in CO2 and H2O carry less energy than the covalent bonds in
wood. The energy released by the initial burning wood promotes the
breaking of more covalent bonds in the wood, and the fire spreads
while more CO2 and H2O are produced. Smoke, ash, and wasted heat
are examples of increased entropy, abbreviated S, as a consequence of
burning the wood. Chemists and physicists have quantified the intercon-
version of energy, heat, and work in three components:

Total energy of the reaction = H (total release of heat from

the fire)
Free energy available to do work = G (heat absorbed by pot
and water)
Entropy or disorder = S (unused energy in smoke, ashes and
heated air)

When looking at the wood before it burned, there was a lot of poten-
tial energy. The burned wood contained much less energy than the
unburned wood. Converting unburned wood to burned wood released
the energy of the system (H). The change in H (H = change in the heat
of the reaction; Greek letter delta () indicates change) value for burning
wood would be negative, meaning the ashes and smoke (H2O and CO2)
contain less H than the unburned wood did. It makes sense, therefore,
that the G for burning wood is also negative, because G is a subset of
H. Using the first law of thermodynamics, all of the heat stored in wood
 Molecules Carry Energy in their Covalent Bonds 3

has to be somewhere, and this law of nature can be described quantitatively

as: G = H S T which looks like a game to guess the word g-h-o-
s-t. This equation says that the change () in energy available to do work
(G = free energy used to boil water) is the difference between the change
in heat of the reaction (H = change in the woods potential energy) and
the change in disorder (S = ashes, smoke, and heat lost to the area) times
the temperature (T = air temperature before the fire started). Burning
wood produces S with a positive value, but G and H are both nega-
tive values. Covalent bonds in wood that contain larger amounts of free
energy are less stable, and they break more easily than the more stable,
lower energy covalent bonds in CO2 and H2O. The air temperature in the
entire campsite is unchanged (due to its vast size), as is the air pressure, so
these components of the equation are constant while the fire is burning.
It might be difficult to picture where woods energy is located prior
to its burning. To help see the source of woods hidden energy, think of
woods molecular structure as a series of atoms held together by covalent
bonds (Figure 1). ATP is a high energy molecule that should be famil-
iar from previous studies in biology. ATP is a nucleotide and an energy
source, but where is the energy? Every covalent bond contains a pair of

covalent bond

phosphorous

nitrogen

oxygen

carbon
hydrogen

Figure 1 Covalent bonds contain high energy electrons. A molecule

of ATP with atoms depicted as spheres and covalent bonds depicted
as thick lines connecting atoms of carbon, oxygen, hydrogen, nitrogen
and phosphorous.
Source: Original art from ATP Jsmol.
4 CELLULAR RESPIRATION

shared electrons, and the bond that will be consumed in most ATP re-
actions is the bond that holds the last phosphate onto the penultimate
phosphate. Remember that electricity is moving electrons, and the elec-
trons in the covalent bonds of wood are also a source of energy. Wood
contains trillions of covalent bonds, and their energy is released as heat
when fire breaks the covalent bonds. Chapter 2 will expand on the release
of energy from covalent bonds. Until then, it is important to understand
how chemists can measure the amount of energy in bonds.
Chemists have devised two different pieces of equipment to measure the
heat of a reaction H and the amount of energy available to do work G
(Figure 2). The bomb calorimeter is a modification of the first machines
designed in the 1780s to measure the amount of heat energy given off from
breaking all the covalent bonds of a compound. The modern versions of
these devices are essentially unchanged and consist of five basic parts: a water
bath, a reaction chamber, a stirring rod, a thermometer, and a spark gen-
erator to initiate the chemical reaction. The reaction chamber is filled with
the compound of interest and oxygen gas to facilitate the burning. As the

thermometer
(measure = heat
stir of the reaction, H)
voltameter
spark 0
reactant 1 reactant 2
V +V
e e
e e
water e e
R1ox R2ox
oxygen R1red R2red
compound +
+
+ + + +
A B

Figure 2 Measuring energy released from chemical bonds. A, Bomb

calorimeter burns a compound of interest (H) inside a container
submerged in water. B, Electrochemical cell allows electrons to flow
to reduce reactant 2, while counter ions flow to oxidize reactant 1.
The device measures voltage, and you calculate G from voltage.
Source: Original art.
 Molecules Carry Energy in their Covalent Bonds 5

compound loses the energy in its covalent bonds due to burning, the heat is
absorbed by a known volume of water, and the increase in water temperature
is quantified. The heat is derived from the chemical breakdown of covalent
bonds the same way wood releases its energy when burned. The amount of
releasable heat in a compound, H, is quantified by a bomb calorimeter.
Biologists tend to be more interested in the amount of free energy
available to do molecular work, G (heat used to boil the water) rather
than the total released heat of the reaction, H, which includes smoke,
ashes and wasted heat. To understand what G is, it is important to
understand how G is determined experimentally in an electrochemical
cell (Figure 2B). By tradition, the reaction container on the left is used
as a standard chemical reaction, such as the interconversion of oxidized
H+ ions and reduced H2 gas. Into the reaction container on the right
is place the compound that is being studied in reduced and oxidized
forms, such as the biologically significant molecules of oxidized nico-
tinamide adenine dinucleotide (NAD+) and reduced nicotinamide ad-
enine dinucleotide plus hydrogen (NADH). Remember from previous
chemistry lessons that reduction is the acquisition of electrons, often
in the form of new covalent bonds. Furthermore, the reaction takes
place in water (the universal solvent) because the molecular polarity
of its partially charged oxygen and hydrogen atoms that allow it to dis-
solve more substances than any other liquid. Oxidation is the loss of
electrons from broken covalent bonds. The standard reaction between
H+ and H2 can serve as an electron source to reduce NAD+ to NADH
in the right chamber of Figure 2B. Alternatively, the compound being
studied might produce an oxidizing reaction and generate electrons
that would reduce H+ to H2. Whichever way the electrons flow, the
voltameter quantifies the rate of electron flow as well as the direction
of the flow, left to right in Figure 2B. A reducing chemical reaction on
the right would produce a positive voltage, whereas an oxidizing reac-
tion would produce a negative voltage. Below the reaction chambers is a
tube that allows positive or negative ions to flow in opposite directions
to keep the overall charge neutral, or maintain chemical homeostasis. In
short, an electrochemical cell exhibits atomic homeostasis because as
one chamber accumulates electrons, it also gets rid of other charged ions
so that the overall chamber remains electrically balanced. The energy of
6 CELLULAR RESPIRATION

the paired electrons in a covalent bond is transferred from the oxidized

reactant to the reduced reactant in the electrochemical cell.
From a biological standpoint, the most important aspect of an elec-
trochemical cell is that it experimentally quantifies the electron flow from
a compound of interest, such as NADH being oxidized to NAD+. The
flow of electrons is electricity and an indirect measure of the amount
of free energy available to perform molecular work, or G. To calculate
the precise value for the change in free energy of a compound as it was
oxidized, G, one plugs the measured voltage from the electrochemical
cell into this equation: G = 2nF; = measured voltage from electro-
chemical cell; F = known constant based on the charge of an electron,
and n = known number of electrons per mole of compound. Look at the
following example for the oxidation of NADH to NAD+:

= voltameter of the overall reaction would read: +1.55 volts

(volts = joules/coulomb)
n = number of electrons per reaction: 2 moles of e- for one
mole of reaction
F = 96,485 coulombs of energy for one mole e- (a constant value)

Look at the units in the equation and simplify the units for G:

joules mole e- coulomb

G = 3 3
coulomb mole reaction mole e-

joules mole e- coulomb

G = 3 3
coulomb mole reaction mole e-

joules
G =
mole reaction

The change in free energy available to perform work, G, quanti-

fies the loss of potential energy if the molecule is being oxidized (burn-
ing wood or consuming ATP), which produced a negative value for G
(Figure 3). ATP is a key unit of energy used to maintain the metabolic
economy of cells and molecules. When ATP loses its terminal phosphate
by the breaking of a covalent bond, adenosine diphosphate (ADP) con-
tains less total potential energy (H) and less free energy available to do
work (G). Referring back to the equation G = H S T that re-
lates G to H, some logical conclusions can be drawn. Remember that
 Molecules Carry Energy in their Covalent Bonds 7

ATP

ADP

+ Pi
G = 32 kJ/mole

Figure 3 Converting ATP to ADP releases energy.

The terminal phosphate (Pi) is removed from ATP and
produced ADP. The products have less energy due to
the two electrons lost in the broken bond.
Source: Original art.

the total energy of a system (H) is greater than the free energy available
to do work (G). Starting with one molecule of ATP and ending with two
molecules (ADP and inorganic phosphate, Pi) in addition to lost heat, the
randomness of the system has increased; S is a positive value. However,
reversing the chemical reaction and investing energy to make ATP from
ADP and Pi, there would be a new covalent bond, increased values for G
(>0) and H (>0), but decreased value for S (<0).
Like steam, shared electrons in covalent bonds are potential energy.
Electrons are negatively charged and thus repel each other. Once liber-
ated from the covalent bond, the electrons carry with them some of the
energy (G = free energy), but other portions of the energy holding the
electrons in the covalent bond will be lost to the environment as heat
(part of the H, or total heat of the reaction). Any time a larger molecule
(such as, ATP) is broken down into two smaller molecules (such as, ADP
and Pi), molecular randomness or disorder (S = entropy) is increased.
When bigger molecules are broken into smaller molecules with increased
disorder, air near the reaction will absorb some vibrational energy, which
we detect as heat as when wood is burned. Therefore, products from
burning wood or consuming ATP have negative values for H and G,
8 CELLULAR RESPIRATION

Table 1 Common biological molecules used in cellular respiration,

and their G values.
reaction G (kJ/mole)
ATP ADP + Pi 32
+ +
NADH + O2 + H NAD + H2O 218
O2 + H2 H2O 236
+ -
pyruvate: H + 5/2O2 + C3H3O3 2H2O + 3CO2 1145
FADH2 + O2 FAD + H2O 199
glucose: 6O2 + C6H12O6 6 H2O + 6CO2 2884

but positive values for S. The production of polymers would produce

positive H and G values but a negative S value, because energy was
invested in new covalent bonds between atoms, which decreased the mo-
lecular disorder in the system. Table 1 gives the G values for some com-
mon biological molecules to increase understanding of where potential
energy is stored.
Think of a covalent bond as a tether holding two atoms together to
understand why some covalent bonds carry more energy than others.
Would two stallions tied together and two mice tied together need teth-
ers of equal strength? Of course not, because antagonistic horses run-
ning away from each other would require a stronger tether compared to
a pair of angry mice. Similarly, some atoms have greater repulsive forces
and thus require stronger covalent bonds and thus electrons with more
energy, both free energy (G) and total energy (H). Bomb calorimeters
measure all of the energy in bonds, which is measured as a change of heat
in the water (H). Electrochemical cells measure the power of electrons
released, which could be used to perform molecular work (G). From
Table 1 it is clear that burning glucose in the presence of O2 to produce
CO2 + H2O released the largest amount of free energy electrons (largest
negative G value), whereas converting ATP to ADP + Pi released the
least amount of free energy (smallest negative G value).
This chapter was intended as a refresher of chemical and physical ter-
minology and the important concept of moving energy from one covalent
bond to another with lower G and H. Cells require energy to main-
tain homeostasis. Each chemical reaction loses some energy to entropy;
 Molecules Carry Energy in their Covalent Bonds 9

so without an investment of more energy, the breakdown of food to ATP

gradually reduces the amount of G and H at each step. The molecules
inside cells have evolved to harness the energy carried by the electrons
and store it for other uses. Chapter 2 presents the main concepts and
highlights of cellular respiration, but all of the steps that take place will
not be presented.

Bibliography
Alberty RA. Calculation of standard transformed Gibbs energies and
standard transformed enthalpies of biochemical reactants. Arch
Biochem Biophys 353(1):116130, 1998.
Atkins P, de Paula J. Physical chemistry for the life sciences. New York
2006, Freeman.
Borsook H, Winegarden HM. On the free energy of glucose and of tripal-
mitin. Proc Natl Acad Sci 16(9):559573, 1930.
Darling S. Computation of the free energy of a-ketoglutarate and
pyruvate from constants of the transamination process. Nature 160:
403404, 1947.
Gibbs JW. A method of geometrical representation of the thermodynamic
properties of substances by means of surfaces. Trans Conn Acad Arts
Sci, 2:382404, 1873.
Nelson D, Cox M. Lehninger principles of biochemistry. ed 3, New York,
2000, Worth Publishing.
 Index
Acetyl-CoA, 16, 20, 26
Allosterically modulated, 2425
Amino acids, deamination of, 2325
ATP
conversion to ADP, 67
production, from digested foods,
4758
synthase, 5254
Benzoic acid, oxidation of, 13, 14
Beta carbon, 1415
Beta oxidation, 1416, 20
biochemical requirements for, 15
Blank acid, 39
Bomb calorimeter, 4, 5, 8
Carbohydrate metabolism, 2532
2-carbon intermediates, energy
extraction, 3946
Casein, 11, 12
Cellular respiration, 19, 20, 25
efficiency calculation of, 56
homeostasis of, 55
substrates for, 5455
Chemical bonds
energy released from,
measuring,45
Chemiosmosis, 52
Cinnamic acid, oxidation of, 14
Citrate, 30
Citric acid cycle, 4344, 45, 47
discovery of, 43
homeostatic regulation of, 44
Coenzyme A (CoA), 16, 1920
Coenzymes, 12
Cofactors, 1213
Covalent bonds, molecular energy
in,19
Deamination of amino acids, 2325
Delineation, 41
Digested foods, ATP production
from, 4758
energy homeostasis, 5457
ethical, legal, social implications
of,5758
Eating vegetables, importance of,
5758
Electricity, 1, 4
Electron transfer pathway,
in mitochondrial
membrane,4951
Energy homeostasis, 5457
Entropy, 2
Enzymes, 12, 57
Exons, 18
Fatty acids, oxidation of, 13, 14, 17
First law of thermodynamics, 12
Flavin adenine dinucleotide
(FAD),13
Foods into energy, converting, 1135
carbohydrate metabolism, 2532
ethical, legal, social implications
of,3235
lipid metabolism, 1320
protein metabolism, 2025
Free energy, 2
Glutamate, 2223
Glutamate dehydrogenase (GDH),
22, 23, 2425
Glycolysis, 26, 27
homeostatic regulation of, 2930
GTP, 2425, 39, 41, 45, 47, 51, 54,
56, 61
Homeostasis, 5
Homozygous, 18
Introns, 18
66 INDEX

Kinase, 30 OMIM, 17
Krebs, Hans Adolf, 39, 40, 42 Oxaloacetic acid, 3943
Oxidation, 5, 14
Lactase, 25 beta, 1416, 20
Lactose, 11, 12 of fatty acids, 13, 14, 17
Lehninger, Albert, 1617, 4344, 47
Limiting factor, 20 Palmitic acid, 11, 12
Lipid metabolism, 1320 degradation pathway for, 17
diseases, genetic basis for, 18 Pedigree, 18
limiting factor in, 20 Peptide bonds, 20
Lipids, 20 Phenylacetic acid, oxidation
Long chain acetyl-CoA dehydrogenase of, 13, 14
(LCAD), 19 Phenylbutyric acid, oxidation of, 14
Phenyl isocrontonic acid,
Malnutrition, 58 oxidationof, 14
Mayer, Richard, 33 Phenylproprionic acid, oxidation
Messenger RNA (mRNA), 19 of,14
Micronutrients, 57 Phosphofructokinase (PFK), 2931
Mitchell, Peter, 5152 Protease, 2122
Mitochondrial matrix, 17, 47 Protein metabolism, 2025
Mitochondrial membrane, electron Pyruvate, 26
transfer pathway in, 4951
Molecular energy, in covalent Quick, Armand, 1314
bonds,19
Multicellular organisms, 56 Redox, 12
Reduced, 5
Negative feedback loop, 30
Nicotinamide adenine dinucleotide Second law of thermodynamics, 2
(NAD+), 5, 13, 24
NADH to, oxidation of, 6 Thermodynamics, 1
Nicotinamide adenine dinucleotide first law of, 12
plus hydrogen (NADH), 5, second law of, 2
22, 24, 25, 61
to NAD+, oxidation of, 6 Very long chain acetyl-CoA
Nutrient deficiencies, 58 dehydrogenase (VLCAD),
1718, 19
Octanoic acid, beta oxidation of, 15
Olestra, 15 Work, 1
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