CHAPTER 13

HOW CELLS OBTAIN ENERGY FROM FOOD

2004 Garland Science Publishing

The Breakdown of Sugars and Fats

13-1

Which of the following stages in the breakdown of the piece of toast you had for
breakfast generates the most ATP?
(a)
Digestion of starch to glucose
(b)
Glycolysis
(c)
The citric acid cycle
(d)
Oxidative phosphorylation
(e)
Conversion of pyruvate to acetyl CoA

13-2

The advantage to the cell of the gradual oxidation of glucose during cellular respiration
compared with its combustion to CO2 and H2O in a single step is
(a)
more free energy is released for a given amount of glucose oxidized.
(b)
no energy is lost as heat.
(c)
energy can be extracted in usable amounts.
(d)
more CO2 is produced for a given amount of glucose oxidized.
(e)
less O2 is required for a given amount of glucose oxidized.

13-3

The final metabolite produced by glycolysis is

(a)
acetyl CoA.
(b)
pyruvate.
(c)
3-phosphoglycerate.
(d)
glyceraldehyde 3-phosphate.
(e)
fatty acids.

13-4

Which of the following steps or processes in aerobic respiration include the production of
carbon dioxide?
(a)
Breakdown of glycogen
(b)
Glycolysis
(c)
Conversion of pyruvate to acetyl CoA
(d)
Oxidative phosphorylation
(e)
The citric acid cycle

207

13-5

On a diet consisting of nothing but protein, which of the following is the most likely
outcome?
(a)
Loss of weight because amino acids cannot be used for the synthesis of fat.
(b)
Muscle gain because the amino acids will go directly into building muscle.
(c)
Tiredness because amino acids cannot be used to generate energy.
(d)
Excretion of more nitrogenous (ammonia-derived) wastes than with a more
balanced diet.
(e)
Production of more carbon dioxide than with a more balanced diet.

13-6

Figure Q13-6 represents a cell lining the gut. Draw numbered labeled lines to indicate
exactly where inside a cell the following processes take place.

Figure Q13-6
1.
2.
3.
4.
5.
6.
7.

Glycolysis
Citric acid cycle
Conversion of pyruvate to activated acetyl groups
Oxidation of fatty acids to acetyl CoA
Glycogen breakdown
Release of fatty acids from triacylglycerols
Oxidative phosphorylation

13-7

Each of the ten steps of glycolysis is catalyzed by a different enzyme. Steps 1 and 3,
catalyzed by hexokinase and phosphofructokinase, involve the hydrolysis of one ATP
molecule. Steps 7 and 10, catalyzed by phosphoglycerate kinase and pyruvate kinase,
each generates a single ATP molecule. At first glance, it seems that the final ATP yield is
zero because there are two ATP-hydrolysis steps and two ATP-formation steps. How can
the net yield of glycolysis be two ATP molecules per glucose molecule?

13-8

The oxidation of sugars by glycolysis

(a)
occurs only in aerobic organisms.
(b)
generates carbon dioxide.
(c)
produces a net gain of ATP.
(d)
occurs in mitochondria.
(e)
uses NADH as a source of energy.

208

13-9

Phosphorylation of glucose (GLC) to produce glucose 6-phosphate (G6P) is the first step
in glucose metabolism after entry into cells. Thermodynamically, it is perfectly valid to
consider the cellular phosphorylation of glucose as the sum of two reactions.

NET:

GLC + Pi
ATP + H2O
GLC + ATP

G6P + H2O
ADP + Pi
G6P + ADP

G = 3.3 kcal/mole
G = 7.3 kcal/mole

But biologically it makes no sense at all. Hydrolysis of ATP (reaction 2) in one part of
the cell can have no effect on phosphorylation of glucose (reaction 1) elsewhere in the
cell.
A.
How does the cell manage to link these two reactions?
B.
What is the G for the net reaction?
13-10 What purpose is served by the phosphorylation of glucose to glucose 6-phosphate by the
enzyme hexokinase as the first step in glycolysis?
(a)
It helps drive the uptake of glucose from outside the cell.
(b)
It generates a high-energy phosphate bond.
(c)
It converts ATP to a more useful form.
(d)
It enables the glucose 6-phosphate to be recognized by phosphofructokinase, the
next enzyme in the glycolytic pathway.
(e)
It oxidizes one of the carbon atoms to yield usable energy.
13-11 Which reaction does the enzyme phosphoglucose isomerase catalyze?
(a)
glucose glucose 6-phosphate
(b)
fructose 6-phosphate fructose 1,6-bisphosphate
(c)
glucose 6-phosphate fructose 6-phosphate
(d)
glucose glucose 1-phosphate
(e)
glucose fructose
13-12 Give the full names of the reactants indicated by question marks in Figure Q13-12.

Figure Q13-12

209

13-13 For each statement below, indicate whether it is TRUE or FALSE. Explain why.
A.
Only aerobic organisms can do glycolysis, suggesting that glycolysis evolved
rather recently.
B.
Oxidation of a molecule requires the removal of electrons and can occur even if
there is no oxygen involved in the reaction.
C.
For a cell to capture energy from oxidation of food molecules, it is better to
release the energy in small packets so it can be stored in activated carrier
molecules.
D.
Fermentation produces more ATP than glycolysis.
E.
One turn of the citric acid cycle generates two molecules of CO2.
F.
The breakdown of one molecule of glucose during glycolysis results in one
molecule of pyruvate.
G.
NADH is more reduced than NAD+.
H.
The reactions of the citric acid cycle do not directly require the presence of
oxygen.
13-14 Which of the following cells rely exclusively on glycolysis to supply them with ATP?
(a)
Anaerobically growing yeast.
(b)
Aerobic bacteria.
(c)
Skeletal muscle cells.
(d)
Plant cells.
(e)
Protozoa.
13-15 In anaerobic conditions, skeletal muscle produces
(a)
lactate and CO2.
(b)
ethanol and CO2.
(c)
lactate only.
(d)
ethanol only.
(e)
lactate, ethanol, and CO2.
13-16 In mammals, liver cells are able to convert lactate to pyruvate. What purpose does this
serve for the organism?
(a)
It is an important way of generating more NADH for the organism.
(b)
It is an important way of generating NAD+.
(c)
It allows the organism to grow in anaerobic conditions.
(d)
It allows the lactate to be productively utilized.
(e)
It is an important way for the body to generate heat.
13-17 Anaerobically growing yeast further metabolizes the pyruvate produced by glycolysis to
CO2 and ethanol as part of a series of fermentation reactions.
A.
What other important reaction occurs during this fermentation step?
B.
Why is this reaction (i.e., the answer to part A) essential for the anaerobically
growing cell?

210

13-18 In the absence of oxygen, cells consume glucose at a high, steady rate. When oxygen is
added, glucose consumption drops precipitously and is then maintained at the lower rate.
Why is glucose consumed at a high rate in the absence of oxygen and at a low rate in its
presence?
13-19 Glycolysis and the citric acid cycle are comprised of several types of reactions that occur
sequentially and serve to harvest energy from the oxidation of carbon atoms. From the
two lists below, match the general class of enzyme from list 1 with the type of reaction
catalyzed from list 2.
List 1
A. Kinase
B. Isomerase
C. Dehydrogenase
D. Synthase or synthetase

List 2
1. Generation of product with the same chemical
formula as the substrate but different connections
between atoms.
2. Transfer of phosphate group from one molecule to
another.
3. Formation of additional carbon-carbon bonds.
4. Oxidation of a substrate.

13-20 The first energy-generating steps in glycolysis begin when glyceraldehyde 3-phosphate
undergoes an energetically favorable reaction in which it is simultaneously oxidized and
phosphorylated by the enzyme glyceraldehyde 3-phosphate dehydrogenase to form 1,3bisphosphoglycerate, with the accompanying conversion of NAD+ to NADH. In a second
energetically favorable reaction catalyzed by a second enzyme, the 1,3bisphosphoglycerate is then converted to 3-phosphoglycerate, with the accompanying
conversion of ADP to ATP. Which of the following statements is TRUE?
(a)
The reaction glyceraldehyde 3-phosphate 1,3-bisphosphoglycerate should be
inhibited when levels of NADH fall.
(b)
The G for the oxidation of the aldehyde group on glyceraldehyde 3-phosphate
to form a carboxylic acid is more negative than the G for ATP hydrolysis.
(c)
The high-energy bond to the phosphate group in glyceraldehyde 3-phosphate
contributes to driving the reaction forward.
(d)
The cysteine side chain on the enzyme is oxidized by NAD+.
(e)
The overall reaction glyceraldehyde 3-phosphate 3-phosphoglycerate has a
positive G.

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13-21 The simultaneous oxidation and phosphorylation of glyceraldehyde 3-phosphate

described in Question 13-20 involves the formation of a highly reactive covalent thioester
bond between a cysteine side chain (reactive group SH) on the enzyme (glyceraldehyde
3-phosphate dehydrogenase) and the oxidized intermediate (see arrow in Figure Q1321A). If the enzyme had a serine (reactive group OH) instead of a cysteine at this
position, which could form only a much lower-energy bond to the oxidized substrate (see
arrow in Figure Q13-21B), how might this new enzyme act?

Figure Q13-21
(a)
(b)
(c)
(d)
(e)

It would oxidize the substrate and phosphorylate it without releasing it.

It would oxidize the substrate but not release it.
It would phosphorylate the substrate on the 2 position instead of the 1 position.
It would behave just like the normal enzyme.
It would use ATP instead of Pi to phosphorylate the substrate.

13-22 Acetyl CoA is

(a)
synthesized from pyruvate in the mitochondrial intermembrane space.
(b)
the intermediate through which food molecules are completely metabolized to
carbon dioxide in animal cells.
(c)
synthesized from pyruvate and CoA in a reaction that also generates NADH, CO2,
and ATP.
(d)
synthesized by the breakdown of fatty acids in the cytosol.
(e)
an intermediate in the oxidation of glucose in anaerobic skeletal muscle.

212

13-23 Assuming complete oxidation, which of the fatty acids shown in Figure Q13-23 will
generate the most ATP through cellular respiration? Why?

Figure Q13-23
13-24 During a single turn of the citric acid cycle
(a)
the two carbon atoms from acetyl CoA that enter the cycle are completely
oxidized to CO2.
(b)
three molecules of ATP are generated.
(c)
three molecules of NADH are generated.
(d)
an acetyl group is added to citric acid.
(e)
three molecules of CO2 are generated.
13-25 Explain why the following statement is FALSE.
One mole of oxaloacetate is required for every mole of acetyl CoA that is
metabolized via the citric acid cycle.
13-26 Cells oxidizing acetyl groups via the citric acid cycle require molecular oxygen in order
to
(a)
oxidize the acetyl groups to CO2.
(b)
regenerate NAD+.
(c)
regenerate FADH2.
(d)
regenerate CoA.
(e)
oxidize fatty acids to acetyl groups.
13-27 Given a mixture of all the enzymes of the citric acid cycle plus acetyl CoA, which of the
following sets of additions could support conversion of acetyl CoA to carbon dioxide?
Explain why.
(a)
Water, NAD+, GDP, phosphate, FAD+
(b)
Water, NAD+, GDP, phosphate, FAD+, oxaloacetate
(c)
Water, NAD+, GDP, phosphate, FAD+, citrate
(d)
Water, NAD+, GDP, phosphate, FAD+, citrate, coenzyme A
13-28 The last reaction of the citric acid cycle, which regenerates oxaloacetate (OAA) from
malate (MAL), has a very positive G = 7.1 kcal/mole. Despite its unfavorable
equilibrium position, material must flow through this reaction quite readily in
mitochondriaotherwise the cycle could not turn. How is flow through the cycle
accomplished in the face of such an overwhelmingly positive G?
213

13-29 Consider the schematic of the citric acid cycle shown in Figure Q13-29. The cycle
begins with the formation of a 6-carbon (6C) molecule from joining the 4-carbon (4C)
oxaloacetate and the 2-carbon (2C) acetyl group. By about halfway through the cycle,
two carbons have been lost as carbon dioxide and a 4C molecule labeled D is
regenerated. List two reasons why the cycle does not stop after production of D and
instead must continue through a series of different 4C intermediates.

Figure Q13-29
13-30 Which of the following statements regarding electron transport is TRUE?
(a)
Only high-energy electrons from NADH can be used to drive the electron
transport chain.
(b)
The proteins involved in electron transport couple oxidation to phosphorylation in
much the same way that glyceraldehyde 3-phosphate dehydrogenase couples
oxidation to phosphorylation in glycolysis.
(c)
Electron transport occurs only in eucaryotes.
(d)
Molecular oxygen is required as a donor of electrons to the electron transport
chain.
(e)
Electrons passing along the electron transport chain move to successively lower
energy states.
13-31 In the final stage of the oxidation of food molecules, a gradient of protons is formed
across the inner mitochondrial membrane, which is normally impermeable to protons. If
cells were exposed to an agent that causes the membrane to become freely permeable to
protons, which of the following effects would you expect to observe?
(a)
Cells would be completely unable to synthesize ATP.
(b)
NADH would build up.
(c)
Carbon dioxide production would cease.
(d)
Consumption of oxygen would fall.
(e)
The ratio of ATP to ADP in the cytoplasm would fall.

214

13-32 For each of the following sentences, fill in the blanks with the best word or phrase
selected from the list below. Not all words or phrases will be used; each word or phrase
may be used more than once.
Oxidative phosphorylation is a process that occurs in the
__________________ of mitochondria. It requires an electron-transport
chain that operates on the high-energy electrons taken from the activated
carrier molecules __________________ and __________________ that
are produced by glycolysis and the citric acid cycle. These electrons are
transferred through a series of molecules, and the energy released during
these transfers is used to generate a gradient of __________________, or
__________________. Since their concentration is much
__________________ outside than inside the mitochondria, the flow of
__________________, or __________________, down the concentration
gradient is energetically very __________________ and can thus be
coupled to the production of ATP from ADP. Thus, oxidative
phosphorylation refers to the oxidation of __________________ and
__________________ molecules and the phosphorylation of
__________________. Without this process, the yield of ATP from each
glucose molecule would be __________________ decreased.
ADP
ATP
cytosol
electrons
FADH2
favorable
glucose

GTP
H+
higher
inner membrane
lower
matrix
moderately

NAD+
NADH
Pi
protons
severely
slightly
unfavorable

How We Know: Unraveling the Citric Acid Cycle

13-33 Consider the following statement: Oxaloacetate acts catalytically to aid in the oxidation
of pyruvate found in suspensions of minced pigeon muscles.
A.
What was the original evidence for this statement?
B.
How did these sorts of experiments aid in the identification of intermediates in the
citric acid cycle?

215

Storing and Utilizing Food

13-34 Which of the following statements is TRUE?
(a)
Plant cells store all their food reserves as starch, whereas animals store all their
food reserves as glycogen.
(b)
Glycogen stores more energy than starch because glycogen molecules have many
more branch points that can be hydrolyzed.
(c)
Animal cells can convert fatty acids to sugars.
(d)
Plants synthesize starch for the same reason that animals synthesize glycogen.
(e)
Protein is an important form of energy storage in animal cells under normal
conditions.
13-35 In humans, glycogen is a more useful food storage molecule than fat because
(a)
a gram of glycogen produces more energy than a gram of fat.
(b)
it can be utilized to produce ATP under anaerobic conditions whereas fat cannot.
(c)
it binds water and therefore is useful in keeping the body hydrated.
(d)
for the same amount of energy storage, glycogen occupies less space in a cell than
does fat.
(e)
glycogen can be carried to cells via the bloodstream whereas fats cannot.
13-36 The intermediates of the citric acid cycle are constantly being depleted because they are
used to produce many of the amino acids needed to make proteins. These intermediates
must therefore be replenished by the conversion of pyruvate to oxaloacetate by the
enzyme pyruvate carboxylase. Bacteria, but not animal cells, have additional enzymes
that can carry out the reaction acetyl CoA + isocitrate oxaloacetate + succinate. Which
of the following compounds will not support the growth of animal cells when used as the
major source of carbon in food, but will support the growth of nonphotosynthetic
bacteria?
(a)
Pyruvate
(b)
Glucose
(c)
Fatty acids
(d)
Carbon dioxide
(e)
Fructose
13-37 Pyruvate can be converted into many other molecules by various biosynthetic and
metabolic pathways, which makes it a central hub in the regulation of cellular
metabolism. Which of the following molecules is not made from pyruvate?
(a)
Oxaloacetate
(b)
Ethanol
(c)
NADH
(d)
Lactate
(e)
Acetyl CoA

216

13-38 For each of the following sentences, fill in the blanks with the best word or phrase
selected from the list below. Not all words or phrases will be used; each word or phrase
should be used only once.
A carbon atom in a CO2 molecule in the atmosphere eventually becomes a
part of one of the enzymes that catalyzes glycolysis in one of your cells.
The CO2 first enters a cell in a corn leaf where photosynthesis fixes the
carbon to make it part of a sugar molecule, which travels from the leaf to
an ear of corn where it is stored as part of a polysaccharide
__________________ molecule in the corn seed. You then eat a corn chip
made from the corn seed. You digest the corn seed, and the free
__________________ travels in your bloodstream, eventually being taken
up by a liver cell and stored as __________________. When required,
this storage molecule breaks down into glucose-1-phosphate, which enters
the glycolytic pathway. Glycolysis produces __________________,
which is converted into acetyl CoA, which enters the
__________________. Several intermediates in this process can provide
the carbon skeleton for production of __________________, which are
then incorporated into the enzymes that catalyze steps in glycolysis.
amino acids
carbon fixation
citric acid cycle
fatty acid
fermentation
galactose
glucose
glycogen

insulin
lactate
nucleotides
oxidative phosphorylation
pyruvate
starch
triacylglycerol

217

Answers
13-1

Choice (d) is the correct answer.Oxidative phosphorylation produces about 28 ATP

molecules. Choice (a) produces no ATP; choice (b) nets 2 ATP; choice (c) produces 1
GTP; and choice (e) produces no ATP.

13-2

Choice(c) is the correct answer. Choice (a) is untrue as the same overall amount of free
energy is released by glucose oxidation, whatever the route. Choice (b) is untrue as a
proportion of the energy released is still lost as heat. Choices (d) and (e) are untrue as the
same amount of CO2 will be released and O2 consumed by the oxidation of glucose to
CO2 and H2O, whatever the route.

13-3

(b)

13-4

Choices (c) and (e) are the correct answers. To obtain the maximal energy from pyruvate,
all three carbons can be fully oxidized to carbon dioxide. First, one carbon is oxidized
and released as carbon dioxide when pyruvate is converted to acetyl CoA. Second, the
two remaining carbons are fully oxidized and released as carbon dioxide by virtue of the
citric acid cycle.

13-5

Choice (d) is the correct answer. Because ammonia is given off when amino acids are
metabolized to yield energy but is not given off when sugars and fats are metabolized,
you would expect more nitrogenous waste to be excreted. Choice (c) is incorrect since
amino acids can be converted into pyruvate and acetyl CoA and used to generate energy.
If more amino acids are consumed than are used, the body will not store them as protein
in muscle tissue but rather will store them as fat, so choices (a) and (b) are incorrect.
Amino acid metabolism does not produce more carbon dioxide than carbohydrate or fat
metabolism, so choice (e) is incorrect.

13-6

See Figure A13-6.

Insert Figure A13-16

218

13-7

The net yield of glycolysis is two ATP molecules per glucose molecule because the steps
that require ATP occur once per glucose molecule whereas those that generate ATP occur
twice per glucose molecule. This is because Step 4, catalyzed by aldolase, breaks down a
6-carbon intermediate into two 3-carbon intermediates. Each 3-carbon intermediate is
processed by the remaining steps and thus Steps 7 and 10 occur twice per original 6carbon glucose molecule.

13-8

Choice (c) is the correct answer. Glycolysis, the step-wise oxidation of glucose to
pyruvate, produces a net gain of two ATP molecules per starting glucose molecule.
Glycolysis occurs in both anaerobic and aerobic organisms; the citric acid cycle, but not
glycolysis, generates carbon dioxide and occurs in mitochondria (thus choices (a), (b),
and (d) are incorrect). Glycolysis does not use NADH and instead produces it (thus
choice (e) is incorrect).

13-9

A.

B.

The cell links these two reactions by using the same enzyme to catalyze both of
them. In essence, the enzyme hexokinase binds both GLC and ATP substrates and
catalyzes the direct transfer of a phosphate group from ATP onto the GLC.
Reactions 1 and 2 do not actually occur because free Pi is never produced.
The G for the net reaction is 4.0 kcal/mole because the free energy is the sum
of component reactions and is independent of the pathway used to convert
substrates into products. G = 3.3 kcal/mole 7.3 kcal/mole = 4.0 kcal/mole.

13-10 Choice (a) is the correct answer. It helps drive the uptake of glucose from outside the cell.
Choice (b) is incorrect since the phosphate transferred to the glucose is not held by a
high-energy covalent bond. Choice (c) is incorrect since the reaction converts ATP to
ADP, which is not useful as an energy source for most cellular reactions, even though it
still has one high-energy bond. Choice (d) is incorrect since the next enzyme in the
pathway is phosphoglucose isomerase, not phosphofructokinase. Choice (e) is incorrect
since the reaction does not involve oxidation of carbon nor does it yield usable energy.
13-11 Choice (c) is the correct answer. The isomerase part of the enzyme name indicates that it
catalyzes an isomerization reaction, and the phosphoglucose part of the name indicates
the type of substrate used. The enzyme that catalyzed reaction (e) would be called
glucose isomerase. The enzymes that catalyzed reactions (a), (b), and (d) would be called
kinases, because they transfer phosphate groups from one molecule to another.
13-12 phosphoenolpyruvate, adenosine diphosphate, pyruvate, adenosine triphosphate

219

13-13 A.
B.
C.
D.
E.
F.
G.
H.

13-14 (a)

False. All organisms can do glycolysis, suggesting that it evolved rather early.
True. Oxidation is the loss of electrons. The mechanism of oxidation often
involves reaction with oxygen atoms, but instead can occur in other ways.
True. If all the energy of oxidation were released at once, as in combustion, most
of the energy would be lost as heat and not captured in usable forms.
False. Fermentation by itself produces no ATP, but only regenerates NAD+ so that
glycolysis can continue to operate and to produce ATP.
True. Each turn of the citric acid cycle essentially exchanges the two carbons
from the input acetyl group into two carbons completely oxidized as CO2.
False. Glycolysis converts each molecule of glucose into two molecules of
pyruvate.
True. NADH is more reduced than NAD+, because it contains more electrons.
True. Oxygen is not one of the substrates in the reactions of the citric acid cycle.
Instead oxygen is needed for the citric acid cycle only indirectly, to regenerate the
oxidized forms of NAD+ and FAD, which are substrates of the citric acid cycle.
All the other cells can carry out oxidative phosphorylation to generate additional
ATP.

13-15 (c)
13-16 Choice (d) is the correct answer. Lactate is a metabolic dead end and it cannot be utilized
any further by cell. Pyruvate, on the other hand, can either be converted to acetyl CoA
and metabolized in the citric acid cycle or converted to glucose by a process called
gluconeogenesis. Conversion of lactate to pyruvate uses up NAD+ (making choice (b)
incorrect) and generates NADH, but the reason animal cells make lactate in the first place
is that they are suffering from an excess of NADH (and thus choice (a) is incorrect).
Choice (c) is incorrect since no mammal can survive without oxygen. The amount of heat
generated by converting lactate to pyruvate is negligible, so choice (e) is incorrect.
13-17 A.
B.

NADH NAD+.
Under anaerobic conditions, it is the only means of regenerating the NAD+
required for glycolysis, the main energy-generating pathway of an anaerobically
growing yeast cell.

13-18 Glucose is consumed at a much higher rate in the absence of oxygen because less usable
energy can be harvested from glucose in the absence of oxygen. Regardless of the
presence of oxygen in the environment, cells need about the same amount of energy in
the form of ATP. In the absence of oxygen, a glucose molecule yields only 2 ATP
molecules and thus many glucose molecules must be consumed to satisfy the energetic
needs of the cell. In the presence of oxygen, a glucose molecule yields about 30 ATP
molecules.
13-19 A2; B1; C4; D3

220

13-20 Choice (b) is the correct answer. This is another way of stating that the energetically
favorable oxidation of glyceraldehyde 3-phosphate provides sufficient energy to
ultimately drive the energy-requiring step of ATP synthesis from ADP. Choice (a) is
untrue: NADH is an end product of the reaction G-3-P to 1,3-PG and therefore high (not
low) levels of it would inhibit the reaction. Choice (c) is untrue: the reactions do not
involve the 3-phosphate group on glyceraldehyde 3-phosphate at all. Choice (d) is untrue,
since the cysteine on the enzyme is important in making a covalent intermediate with the
substrate and is not oxidized by NAD+. Choice (e) is untrue, since if the reaction had an
overall positive G, it could not be used to power the energetically unfavorable reactions
of ATP and NADH synthesis.
13-21 Choice (b) is the correct answer. The phosphorylation and release of the product from the
normal enzyme is possible because the high-energy thioester bond formed between the
oxidized substrate and enzyme can be attacked by a phosphate molecule. If the bond
between oxidized substrate and enzyme is a much lower-energy bond, the enzyme will
not be able to transfer the oxidized substrate to a phosphate group, and substrate and
enzyme will remain covalently bound. Choices (a), (c), and (d) could not happen, as none
of the bonds in the substrate molecule is reactive enough to be broken by a phosphate
group. Choice (e) would not happen because the enzyme does not bind ATP.
13-22 Choice (b) is the correct answer. Some food molecules can enter the citric acid cycle at
points other than acetyl CoA, but only molecules that enter as acetyl CoA are completely
oxidized to carbon dioxide. Acetyl CoA is made in the mitochondrial matrix, not the
intermembrane space, so choice (a) is incorrect. Choice (c) is incorrect since no ATP is
directly generated by the conversion of pyruvate to acetyl CoA; carbon dioxide and
NADH are the only other products of the reaction. Choice (d) is incorrect as the
breakdown of fatty acids to produce acetyl CoA occurs in the mitochondrial matrix.
Choice (e) is incorrect since acetyl CoA is not an intermediate in the anaerobic
fermentation reaction that coverts pyruvate to lactate.
13-23 (B)

This will produce 2 NADH, 2 FADH2, and 3 acetyl CoA on complete oxidation.
Because of the double bond in A, this fatty acid will produce 2 NADH and 3
acetyl CoA, but only 1 FADH2, since the initial oxidation step using FAD+ that
reduces the two-carbon unit CH2CH2 to CH=CH is not needed for twocarbon units already containing a double bond. So although the amount of acetyl
CoA entering the citric acid cycle will be the same for A and B, fewer reducing
equivalents will eventually enter the electron transport chain from the oxidation of
A, thus less ATP will be produced.

13-24 (c)
13-25 Only small amounts of oxaloacetate are required relative to the amount of acetyl CoA
metabolized because oxaloacetate is regenerated after every round of the citric acid cycle.

221

13-26 Choice (b) is the correct answer. The citric acid cycle generates high-energy electrons
that are passed to NAD+ to form NADH. NADH then donates these electrons to the
electron transport chain that drives oxidative phosphorylation, regenerating the NAD+
needed to keep the citric acid cycle going. The electrons from NADH are passed via the
electron transport chain to oxygen.
13-27

Choices (b) and (d) are correct. To get the cycle turning you need water, NAD+, GDP,
phosphate, FAD, coenzyme A, and at least one intermediate of the citric acid cycle, which
are all provided in choice (d). In addition, choice (b) will be sufficient because
oxaloacetate reacts with acetyl CoA to release a molecule of coenzyme A, which can then
be reused. Choice (c) would produce a small amount of CO2 initially from the added
citrate, but the cycle could not continue since citrate has to go through a step requiring
coenzyme A to complete the cycle. Choice (a) will not work, as this set does not include
any citric acid cycle intermediate.

13-28 The mitochondrion obtains a flow through this reaction by maintaining high
concentrations of substrates and low concentrations of products, so that the G is
negative despite a positive G. The concentration ratio of products to substrates, [OAA]
[NADH]/[MAL][NAD+], must be substantially smaller than the equilibrium constant K to
overcome a positive G value.
13-29 The two reasons are to keep the cycle going and to harvest more energy. (1) The cycle
must regenerate oxaloacetate, which acts catalytically in the citric acid cycle to aid in the
oxidation of many acetyl groups. (2) Additional reactions are required to more fully
oxidize the carbons and harvest energy by producing several activated carrier molecules,
including GTP, FADH2, and NADH.
13-30 Choice (e) is the correct answer. Electrons passing along the electron transport chain
move to successively lower energy states. Choice (a) is untrue as electrons from FADH2
can be used as well. Choice (b) is untrue as the two mechanisms of coupling oxidation to
phosphorylation are quite different. Oxidative phosphorylation involves the oxidation of
NADH to NAD+ by proteins of the electron transport chain. Electron transport then
causes the formation of a proton gradient across a membrane, which drives ATP
synthesis. In contrast, glyceraldehyde 3-phosphate dehydrogenase action involves the
reduction of NAD+ to NADH and uses the G of glyceraldehyde oxidation to form a
high-energy bond that can be attacked directly by a phosphate group. Choice (c) is
untrue, as electron transport occurs in the plasma membrane of procaryotes. Choice (d) is
untrue, as molecular oxygen acts as an acceptor, not a donor, for electrons.

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13-31 Choice (e) is the correct answer. If the inner mitochondrial membrane became permeable
to protons, the electron transport chain would continue to oxidize NADH to NAD+,
transport electrons and pump protons, so the consumption of oxygen would not fall
(choice (d)). However, the energy stored by the protons would be immediately dissipated
as heat when they flowed back across the membrane and thus could not drive the
synthesis of ATP. But, NADH would not build up (choice (b)), and the citric acid cycle
and glycolysis would continue (and thus CO2 would still be produced, contrary to choice
(c)). Since glycolysis and the citric acid cycle produce 2 molecules of ATP and one
molecule of GTP (which can be converted to ATP), respectively, ATP production would
not completely cease (choice (a)), but it would be very much less than normal.
13-32 Oxidative phosphorylation is a process that occurs in the inner membrane of
mitochondria. It requires an electron-transport chain that operates on the high-energy
electrons taken from the activated carrier molecules NADH and FADH2 that are
produced by glycolysis and the citric acid cycle. These electrons are transferred through
a series of molecules, and the energy released during these transfers is used to generate a
gradient of protons, or H+. Since their concentration is much higher outside than inside
the mitochondria, the flow of protons, or H+, down the concentration gradient is
energetically very favorable and can thus be coupled to the production of ATP from ADP.
Thus, oxidative phosphorylation refers to the oxidation of NADH and FADH2 molecules
and the phosphorylation of ADP. Without this process, the yield of ATP from each
glucose molecule would be severely decreased.
13-33 A.

B.

Experiments showed that minced pigeon muscles contained large amounts of

pyruvate, but oxidized this compound rather slowly, so that little oxygen was
consumed and little carbon dioxide was produced. When a tiny amount of
oxaloacetate was added to such muscle preparations, large amounts of oxygen and
carbon dioxide were consumed and produced, respectively. If the added
oxaloacetate was simply being oxidized fully, the oxygen consumption and
carbon dioxide production would be expected to increase only slightly, but instead
the large amount of oxygen consumed suggested that each molecule of added
oxaloacetate aided in the oxidation of many molecules of some other substance.
Analogous experiments showed that the addition of several other compounds, like
succinate and fumarate, had the same consequences as adding oxaloacetate. This
was interpreted as evidence that these compounds are intermediates in the same
pathway and can be converted into, or are derived from, oxaloacetate, which was
later demonstrated more directly.

13-34 Choice (d) is the correct answer. Both starch and glycogen are storage polymers of
glucose. Choice (a) is false, since both plants and animals can also store food as fats and
oils. Choice (b) is false, as although glycogen synthesis requires ATP, no ATP is generated
by its hydrolysis to monomers (so the number of branch points is irrelevant to energy
storage). Choice (c) is false, as animal cells cannot do this. Choice (e) is false, as the use
of protein for energy occurs only under starvation conditions.

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13-35 Choice (b) is the answer. The breakdown of glycogen to glucose does not require oxygen;
the glucose can then enter glycolysis and generate ATP by a fermentation process that
produces lactic acid. In contrast, fats are broken down to acetyl CoA that must enter the
citric acid cycle, which requires oxygen to keep turning. Choice (a) is incorrect, as a
gram of glycogen (wet or dry) produces less energy than a gram of fat. Choice (c) is
incorrect, as the water bound by glycogen is not useful in keeping the body hydrated and
merely contributes to making the glycogen weigh a lot. Choice (d) is incorrect, as the
actual mass of glycogen required to store the same amount of energy is six-fold greater
than the amount of fat. Choice (e) is incorrect, as fats can be carried in the bloodstream. If
the energy stored in glycogen is required by other cells, glycogen is broken down to
glucose, and the glucose is then released into the bloodstream.
13-36 Choice (c) is the correct answer. In oxidative metabolism, fatty acids can only be
converted to acetyl CoA, which is completely oxidized to carbon dioxide through the
citric acid cycle. In addition, bacteria can use some of this acetyl CoA as a source of
carbon atoms to replenish the citric acid cycle, whereas animals cannot. Choices (a), (b),
and (e) are incorrect, since glucose and fructose can be converted to pyruvate, and hence
to citric acid cycle intermediates, in both animal and bacterial cells, while choice (d) is
incorrect, since carbon dioxide cannot be used as a main carbon source by either
nonphotosynthetic bacteria or animal cells.
13-37 (c)

Pyruvate cannot be converted into NADH, but it can be converted into the other
metabolites in one or two steps.

13-38 A carbon atom in a CO2 molecule in the atmosphere eventually becomes a part of one of
the enzymes that catalyzes glycolysis in one of your cells. The CO2 first enters a cell in a
corn leaf where photosynthesis fixes the carbon to make it part of a sugar molecule,
which travels from the leaf to an ear of corn where it is stored as part of a polysaccharide
starch molecule in the corn seed. You then eat a corn chip made from the corn seed.
You digest the corn seed, and the free glucose travels in your bloodstream, eventually
being taken up by a liver cell and stored as glycogen. When required, this storage
molecule breaks down into glucose-1-phosphate, which enters the glycolytic pathway.
Glycolysis produces pyruvate, which is converted into acetyl CoA, which enters the
citric acid cycle. Several intermediates in this process can provide the carbon skeleton
for production of amino acids, which are then incorporated into the enzymes that
catalyze steps in glycolysis.

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