CHAPTER 13

HOW CELLS OBTAIN ENERGY FROM FOOD

2004 Garland Sc!nc! P"#l$%n&
T%! Br!a'd()n (* S"&ar$ and Fa+$
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) !idati"e phosphorylation
(e) #on"ersion of pyru"ate to acetyl #oA
13-2 The ad"antage to the cell of the gradual o!idation of glucose during cellular respiration
compared with its combustion to #$ and %$ in a single step is
(a) more free energy is released for a gi"en amount of glucose o!idi&ed'
(b) no energy is lost as heat'
(c) energy can be e!tracted in usable amounts'
(d) more #$ is produced for a gi"en amount of glucose o!idi&ed'
(e) less $ is re(uired for a gi"en amount of glucose o!idi&ed'
13-3 The final metabolite produced by glycolysis is
(a) acetyl #oA'
(b) pyru"ate'
(c) )*phosphoglycerate'
(d) glyceraldehyde )*phosphate'
(e) fatty acids'
13-4 Which of the following steps or processes in aerobic respiration include the production of
carbon dio!ide?
(a) +reakdown of glycogen
(b) Glycolysis
(c) #on"ersion of pyru"ate to acetyl #oA
(d) !idati"e phosphorylation
(e) The citric acid cycle
20,
13-5 n a diet consisting of nothing but protein, which of the following is the most likely
outcome?
(a) -oss of weight because amino acids cannot be used for the synthesis of fat'
(b) .uscle gain because the amino acids will go directly into building muscle'
(c) Tiredness because amino acids cannot be used to generate energy'
(d) /!cretion of more nitrogenous (ammonia*deri"ed) wastes than with a more
balanced diet'
(e) Production of more carbon dio!ide than with a more balanced diet'
13-6 0igure 12)*3 represents a cell lining the gut' Draw numbered labeled lines to indicate
e!actly where inside a cell the following processes take place'
0igure 12)*3
2' Glycolysis
$' #itric acid cycle
)' #on"ersion of pyru"ate to acti"ated acetyl groups
4' !idation of fatty acids to acetyl #oA
5' Glycogen breakdown
3' 6elease of fatty acids from triacylglycerols
7' !idati"e phosphorylation
13-7 /ach of the ten steps of glycolysis is cataly&ed by a different en&yme' 8teps 2 and ),
cataly&ed by he!okinase and phosphofructokinase, in"ol"e the hydrolysis of one ATP
molecule' 8teps 7 and 29, cataly&ed by phosphoglycerate kinase and pyru"ate kinase,
each generates a single ATP molecule' At first glance, it seems that the final ATP yield is
&ero because there are two ATP*hydrolysis steps and two ATP*formation steps' %ow can
the net yield of glycolysis be two ATP molecules per glucose molecule?
13-8 The o!idation of sugars by glycolysis
(a) occurs only in aerobic organisms'
(b) generates carbon dio!ide'
(c) produces a net gain of ATP'
(d) occurs in mitochondria'
(e) uses :AD% as a source of energy'
20-
13-9 Phosphorylation of glucose (G-#) to produce glucose 3*phosphate (G3P) is the first step
in glucose metabolism after entry into cells' Thermodynamically, it is perfectly "alid to
consider the cellular phosphorylation of glucose as the sum of two reactions'
G-# ; Pi G3P ; %$ G< = )') kcal>mole
ATP ; %$ ADP ; Pi G< = ?7') kcal>mole
:/T@ G-# ; ATP G3P ; ADP
+ut biologically it makes no sense at all' %ydrolysis of ATP (reaction $) in one part of
the cell can ha"e no effect on phosphorylation of glucose (reaction 2) elsewhere in the
cell'
A' %ow does the cell manage to link these two reactions?
+' What is the G< for the net reaction?
13-10 What purpose is ser"ed by the phosphorylation of glucose to glucose 3*phosphate by the
en&yme he!okinase as the first step in glycolysis?
(a) At helps dri"e the uptake of glucose from outside the cell'
(b) At generates a high*energy phosphate bond'
(c) At con"erts ATP to a more useful form'
(d) At enables the glucose 3*phosphate to be recogni&ed by phosphofructokinase, the
ne!t en&yme in the glycolytic pathway'
(e) At o!idi&es one of the carbon atoms to yield usable energy'
13-11 Which reaction does the en&yme phosphoglucose isomerase cataly&e?
(a) glucose glucose 3*phosphate
(b) fructose 3*phosphate fructose 2,3*bisphosphate
(c) glucose 3*phosphate fructose 3*phosphate
(d) glucose glucose 2*phosphate
(e) glucose fructose
13-12 Gi"e the full names of the reactants indicated by (uestion marks in 0igure 12)*2$'
0igure 12)*2$
20.
13-13 0or each statement below, indicate whether it is T6B/ or 0A-8/' /!plain why'
A' nly aerobic organisms can do glycolysis, suggesting that glycolysis e"ol"ed
rather recently'
+' !idation of a molecule re(uires the remo"al of electrons and can occur e"en if
there is no o!ygen in"ol"ed in the reaction'
#' 0or a cell to capture energy from o!idation of food molecules, it is better to
release the energy in small packets so it can be stored in acti"ated carrier
molecules'
D' 0ermentation produces more ATP than glycolysis'
/' ne turn of the citric acid cycle generates two molecules of #$'
0' The breakdown of one molecule of glucose during glycolysis results in one
molecule of pyru"ate'
G' :AD% is more reduced than :AD
;
'
%' The reactions of the citric acid cycle do not directly re(uire the presence of
o!ygen'
13-14 Which of the following cells rely e!clusi"ely on glycolysis to supply them with ATP?
(a) Anaerobically growing yeast'
(b) Aerobic bacteria'
(c) 8keletal muscle cells'
(d) Plant cells'
(e) Proto&oa'
13-15 An anaerobic conditions, skeletal muscle produces
(a) lactate and #$'
(b) ethanol and #$'
(c) lactate only'
(d) ethanol only'
(e) lactate, ethanol, and #$'
13-16 An mammals, li"er cells are able to con"ert lactate to pyru"ate' What purpose does this
ser"e for the organism?
(a) At is an important way of generating more :AD% for the organism'
(b) At is an important way of generating :AD
;
'
(c) At allows the organism to grow in anaerobic conditions'
(d) At allows the lactate to be producti"ely utili&ed'
(e) At is an important way for the body to generate heat'
13-17 Anaerobically growing yeast further metaboli&es the pyru"ate produced by glycolysis to
#$ and ethanol as part of a series of fermentation reactions'
A' What other important reaction occurs during this fermentation step?
+' Why is this reaction (i'e', the answer to part A) essential for the anaerobically
growing cell?
210
13-18 An the absence of o!ygen, cells consume glucose at a high, steady rate' When o!ygen 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 o!ygen and at a low rate in its
presence?
13-19 Glycolysis and the citric acid cycle are comprised of se"eral types of reactions that occur
se(uentially and ser"e to har"est energy from the o!idation of carbon atoms' 0rom the
two lists below, match the general class of en&yme from list 2 with the type of reaction
cataly&ed from list $'
List 1 List 2
A' Cinase 2' Generation of product with the same chemical
formula as the substrate but different connections
between atoms'
+' Asomerase $' Transfer of phosphate group from one molecule to
another'
#' Dehydrogenase )' 0ormation of additional carbon*carbon bonds'
D' 8ynthase or synthetase 4' !idation of a substrate'
13-20 The first energy*generating steps in glycolysis begin when glyceraldehyde )*phosphate
undergoes an energetically fa"orable reaction in which it is simultaneously o!idi&ed and
phosphorylated by the en&yme glyceraldehyde )*phosphate dehydrogenase to form 2,)*
bisphosphoglycerate, with the accompanying con"ersion of :AD
;
to :AD%' An a second
energetically fa"orable reaction cataly&ed by a second en&yme, the 2,)*
bisphosphoglycerate is then con"erted to )*phosphoglycerate, with the accompanying
con"ersion of ADP to ATP' Which of the following statements is T6B/?
(a) The reaction glyceraldehyde )*phosphate 2,)*bisphosphoglycerate should be
inhibited when le"els of :AD% fall'
(b) The GD for the o!idation of the aldehyde group on glyceraldehyde )*phosphate
to form a carbo!ylic acid is more negati"e than the GD for ATP hydrolysis'
(c) The high*energy bond to the phosphate group in glyceraldehyde )*phosphate
contributes to dri"ing the reaction forward'
(d) The cysteine side chain on the en&yme is o!idi&ed by :AD
;
'
(e) The o"erall reaction glyceraldehyde )*phosphate )*phosphoglycerate has a
positi"e G'
211
13-21 The simultaneous o!idation and phosphorylation of glyceraldehyde )*phosphate
described in 1uestion 2)*$9 in"ol"es the formation of a highly reacti"e co"alent thioester
bond between a cysteine side chain (reacti"e group ?8%) on the en&yme (glyceraldehyde
)*phosphate dehydrogenase) and the o!idi&ed intermediate (see arrow in 0igure 12)*
$2A)' Af the en&yme had a serine (reacti"e group ?%) instead of a cysteine at this
position, which could form only a much lower*energy bond to the o!idi&ed substrate (see
arrow in 0igure 12)*$2+), how might this new en&yme act?
0igure 12)*$2
(a) At would o!idi&e the substrate and phosphorylate it without releasing it'
(b) At would o!idi&e the substrate but not release it'
(c) At would phosphorylate the substrate on the $ position instead of the 2 position'
(d) At would beha"e Eust like the normal en&yme'
(e) At would use ATP instead of Pi to phosphorylate the substrate'
13-22 Acetyl #oA is
(a) synthesi&ed from pyru"ate in the mitochondrial intermembrane space'
(b) the intermediate through which food molecules are completely metaboli&ed to
carbon dio!ide in animal cells'
(c) synthesi&ed from pyru"ate and #oA in a reaction that also generates :AD%, #$,
and ATP'
(d) synthesi&ed by the breakdown of fatty acids in the cytosol'
(e) an intermediate in the o!idation of glucose in anaerobic skeletal muscle'
212
13-23 Assuming complete o!idation, which of the fatty acids shown in 0igure 12)*$) will
generate the most ATP through cellular respiration? Why?
0igure 12)*$)
13-24 During a single turn of the citric acid cycle
(a) the two carbon atoms from acetyl #oA that enter the cycle are completely
o!idi&ed to #$'
(b) three molecules of ATP are generated'
(c) three molecules of :AD% are generated'
(d) an acetyl group is added to citric acid'
(e) three molecules of #$

are generated'
13-25 /!plain why the following statement is 0A-8/'
Fne mole of o!aloacetate is re(uired for e"ery mole of acetyl #oA that is
metaboli&ed "ia the citric acid cycle'G
13-26 #ells o!idi&ing acetyl groups "ia the citric acid cycle re(uire molecular o!ygen in order
to
(a) o!idi&e the acetyl groups to #$'
(b) regenerate :AD
;
'
(c) regenerate 0AD%$'
(d) regenerate #oA'
(e) o!idi&e fatty acids to acetyl groups'
13-27 Gi"en a mi!ture of all the en&ymes of the citric acid cycle plus acetyl #oA, which of the
following sets of additions could support con"ersion of acetyl #oA to carbon dio!ide?
/!plain why'
(a) Water, :AD
;
, GDP, phosphate, 0AD
;
(b) Water, :AD
;
, GDP, phosphate, 0AD
;
, o!aloacetate
(c) Water, :AD
;
, GDP, phosphate, 0AD
;
, citrate
(d) Water, :AD
;
, GDP, phosphate, 0AD
;
, citrate, coen&yme A
13-28 The last reaction of the citric acid cycle, which regenerates o!aloacetate (AA) from
malate (.A-), has a "ery positi"e G< = 7'2 kcal>mole' Despite its unfa"orable
e(uilibrium position, material must flow through this reaction (uite readily in
mitochondriaHotherwise the cycle could not turn' %ow is flow through the cycle
accomplished in the face of such an o"erwhelmingly positi"e G<?
213
13-29 #onsider the schematic of the citric acid cycle shown in 0igure 12)*$I' The cycle
begins with the formation of a 3*carbon (3#) molecule from Eoining the 4*carbon (4#)
o!aloacetate and the $*carbon ($#) acetyl group' +y about halfway through the cycle,
two carbons ha"e been lost as carbon dio!ide and a 4# molecule labeled D is
regenerated' -ist two reasons why the cycle does not stop after production of D and
instead must continue through a series of different 4# intermediates'
0igure 12)*$I
13-30 Which of the following statements regarding electron transport is T6B/?
(a) nly high*energy electrons from :AD% can be used to dri"e the electron
transport chain'
(b) The proteins in"ol"ed in electron transport couple o!idation to phosphorylation in
much the same way that glyceraldehyde )*phosphate dehydrogenase couples
o!idation to phosphorylation in glycolysis'
(c) /lectron transport occurs only in eucaryotes'
(d) .olecular o!ygen is re(uired as a donor of electrons to the electron transport
chain'
(e) /lectrons passing along the electron transport chain mo"e to successi"ely lower
energy states'
13-31 An the final stage of the o!idation of food molecules, a gradient of protons is formed
across the inner mitochondrial membrane, which is normally impermeable to protons' Af
cells were e!posed to an agent that causes the membrane to become freely permeable to
protons, which of the following effects would you e!pect to obser"e?
(a) #ells would be completely unable to synthesi&e ATP'
(b) :AD% would build up'
(c) #arbon dio!ide production would cease'
(d) #onsumption of o!ygen would fall'
(e) The ratio of ATP to ADP in the cytoplasm would fall'
214
13-32 0or each of the following sentences, fill in the blanks with the best word or phrase
selected from the list below' :ot all words or phrases will be usedJ each word or phrase
may be used more than once'
!idati"e phosphorylation is a process that occurs in the
KKKKKKKKKKKKKKKKKK of mitochondria' At re(uires an electron*transport
chain that operates on the high*energy electrons taken from the acti"ated
carrier molecules KKKKKKKKKKKKKKKKKK and KKKKKKKKKKKKKKKKKK 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 KKKKKKKKKKKKKKKKKK, or
KKKKKKKKKKKKKKKKKK' 8ince their concentration is much
KKKKKKKKKKKKKKKKKK outside than inside the mitochondria, the flow of
KKKKKKKKKKKKKKKKKK, or KKKKKKKKKKKKKKKKKK, down the concentration
gradient is energetically "ery KKKKKKKKKKKKKKKKKK and can thus be
coupled to the production of ATP from ADP' Thus, o!idati"e
phosphorylation refers to the o!idation of KKKKKKKKKKKKKKKKKK and
KKKKKKKKKKKKKKKKKK molecules and the phosphorylation of
KKKKKKKKKKKKKKKKKK' Without this process, the yield of ATP from each
glucose molecule would be KKKKKKKKKKKKKKKKKK decreased'
ADP GTP :AD;
ATP %; :AD%
cytosol higher Pi
electrons inner membrane protons
0AD%$ lower se"erely
fa"orable matri! slightly
glucose moderately unfa"orable
H() W! /n()0 1nra2!ln& +%! C+rc Acd C3cl!
13-33 #onsider the following statement@ !aloacetate acts catalytically to aid in the o!idation
of pyru"ate found in suspensions of minced pigeon muscles'
A' What was the original e"idence for this statement?
+' %ow did these sorts of e!periments aid in the identification of intermediates in the
citric acid cycle?
214
S+(rn& and 1+l5n& F((d
13-34 Which of the following statements is T6B/?
(a) Plant cells store all their food reser"es as starch, whereas animals store all their
food reser"es as glycogen'
(b) Glycogen stores more energy than starch because glycogen molecules ha"e many
more branch points that can be hydroly&ed'
(c) Animal cells can con"ert fatty acids to sugars'
(d) Plants synthesi&e starch for the same reason that animals synthesi&e glycogen'
(e) Protein is an important form of energy storage in animal cells under normal
conditions'
13-35 An 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 utili&ed 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 "ia 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 con"ersion of pyru"ate to o!aloacetate by the
en&yme pyru"ate carbo!ylase' +acteria, but not animal cells, ha"e additional en&ymes
that can carry out the reaction acetyl #oA ; isocitrate o!aloacetate ; succinate' Which
of the following compounds will not support the growth of animal cells when used as the
maEor source of carbon in food, but will support the growth of nonphotosynthetic
bacteria?
(a) Pyru"ate
(b) Glucose
(c) 0atty acids
(d) #arbon dio!ide
(e) 0ructose
13-37 Pyru"ate can be con"erted into many other molecules by "arious 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 pyru"ate?
(a) !aloacetate
(b) /thanol
(c) :AD%
(d) -actate
(e) Acetyl #oA
216
13-38 0or each of the following sentences, fill in the blanks with the best word or phrase
selected from the list below' :ot all words or phrases will be usedJ each word or phrase
should be used only once'
A carbon atom in a #$ molecule in the atmosphere e"entually becomes a
part of one of the en&ymes that cataly&es glycolysis in one of your cells'
The #$ first enters a cell in a corn leaf where photosynthesis fi!es the
carbon to make it part of a sugar molecule, which tra"els from the leaf to
an ear of corn where it is stored as part of a polysaccharide
KKKKKKKKKKKKKKKKKK molecule in the corn seed' Lou then eat a corn chip
made from the corn seed' Lou digest the corn seed, and the free
KKKKKKKKKKKKKKKKKK tra"els in your bloodstream, e"entually being taken
up by a li"er cell and stored as KKKKKKKKKKKKKKKKKK' When re(uired,
this storage molecule breaks down into glucose*2*phosphate, which enters
the glycolytic pathway' Glycolysis produces KKKKKKKKKKKKKKKKKK,
which is con"erted into acetyl #oA, which enters the
KKKKKKKKKKKKKKKKKK' 8e"eral intermediates in this process can pro"ide
the carbon skeleton for production of KKKKKKKKKKKKKKKKKK, which are
then incorporated into the en&ymes that cataly&e steps in glycolysis'
amino acids insulin
carbon fi!ation lactate
citric acid cycle nucleotides
fatty acid o!idati"e phosphorylation
fermentation pyru"ate
galactose starch
glucose triacylglycerol
glycogen
21,
An$)!r$
13-1 #hoice (d) is the correct answer'!idati"e phosphorylation produces about $M ATP
molecules' #hoice (a) produces no ATPJ choice (b) nets $ ATPJ choice (c) produces 2
GTPJ and choice (e) produces no ATP'
13-2 #hoice(c) is the correct answer' #hoice (a) is untrue as the same o"erall amount of free
energy is released by glucose o!idation, whate"er the route' #hoice (b) is untrue as a
proportion of the energy released is still lost as heat' #hoices (d) and (e) are untrue as the
same amount of #$ will be released and $

consumed by the o!idation of glucose to
#$ and %$, whate"er the route'
13-3 (b)
13-4 #hoices (c) and (e) are the correct answers' To obtain the ma!imal energy from pyru"ate,
all three carbons can be fully o!idi&ed to carbon dio!ide' 0irst, one carbon is o!idi&ed
and released as carbon dio!ide when pyru"ate is con"erted to acetyl #oA' 8econd, the
two remaining carbons are fully o!idi&ed and released as carbon dio!ide by "irtue of the
citric acid cycle'
13-5 #hoice (d) is the correct answer' +ecause ammonia is gi"en off when amino acids are
metaboli&ed to yield energy but is not gi"en off when sugars and fats are metaboli&ed,
you would e!pect more nitrogenous waste to be e!creted' #hoice (c) is incorrect since
amino acids can be con"erted into pyru"ate and acetyl #oA and used to generate energy'
Af 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 dio!ide than carbohydrate or fat
metabolism, so choice (e) is incorrect'
13-6 8ee 0igure A2)*3'
Ansert 0igure A2)*23
21-
13-7 The net yield of glycolysis is two ATP molecules per glucose molecule because the steps
that re(uire ATP occur once per glucose molecule whereas those that generate ATP occur
twice per glucose molecule' This is because 8tep 4, cataly&ed by aldolase, breaks down a
3*carbon intermediate into two )*carbon intermediates' /ach )*carbon intermediate is
processed by the remaining steps and thus 8teps 7 and 29 occur twice per original 3*
carbon glucose molecule'
13-8 #hoice (c) is the correct answer' Glycolysis, the step*wise o!idation of glucose to
pyru"ate, produces a net gain of two ATP molecules per starting glucose molecule'
Glycolysis occurs in both anaerobic and aerobic organismsJ the citric acid cycle, but not
glycolysis, generates carbon dio!ide and occurs in mitochondria (thus choices (a), (b),
and (d) are incorrect)' Glycolysis does not use :AD% and instead produces it (thus
choice (e) is incorrect)'
13-9 A' The cell links these two reactions by using the same en&yme to cataly&e both of
them' An essence, the en&yme he!okinase binds both G-# and ATP substrates and
cataly&es the direct transfer of a phosphate group from ATP onto the G-#'
6eactions 2 and $ do not actually occur because free Pi is ne"er produced'
+' The G< for the net reaction is ?4'9 kcal>mole because the free energy is the sum
of component reactions and is independent of the pathway used to con"ert
substrates into products' G< = )') kcal>mole ? 7') kcal>mole = ?4'9 kcal>mole'
13-10 #hoice (a) is the correct answer' At helps dri"e the uptake of glucose from outside the cell'
#hoice (b) is incorrect since the phosphate transferred to the glucose is not held by a
high*energy co"alent bond' #hoice (c) is incorrect since the reaction con"erts ATP to
ADP, which is not useful as an energy source for most cellular reactions, e"en though it
still has one high*energy bond' #hoice (d) is incorrect since the ne!t en&yme in the
pathway is phosphoglucose isomerase, not phosphofructokinase' #hoice (e) is incorrect
since the reaction does not in"ol"e o!idation of carbon nor does it yield usable energy'
13-11 #hoice (c) is the correct answer' The isomerase part of the en&yme name indicates that it
cataly&es an isomeri&ation reaction, and the phosphoglucose part of the name indicates
the type of substrate used' The en&yme that cataly&ed reaction (e) would be called
glucose isomerase' The en&ymes that cataly&ed reactions (a), (b), and (d) would be called
kinases, because they transfer phosphate groups from one molecule to another'
13-12 phosphoenolpyru"ate, adenosine diphosphate, pyru"ate, adenosine triphosphate
21.
13-13 A' 0alse' All organisms can do glycolysis, suggesting that it e"ol"ed rather early'
+' True' !idation is the loss of electrons' The mechanism of o!idation often
in"ol"es reaction with o!ygen atoms, but instead can occur in other ways'
#' True' Af all the energy of o!idation were released at once, as in combustion, most
of the energy would be lost as heat and not captured in usable forms'
D' 0alse' 0ermentation by itself produces no ATP, but only regenerates :AD
;
so that
glycolysis can continue to operate and to produce ATP'
/' True' /ach turn of the citric acid cycle essentially e!changes the two carbons
from the input acetyl group into two carbons completely o!idi&ed as #$'
0' 0alse' Glycolysis con"erts each molecule of glucose into two molecules of
pyru"ate'
G' True' :AD% is more reduced than :AD
;
, because it contains more electrons'
%' True' !ygen is not one of the substrates in the reactions of the citric acid cycle'
Anstead o!ygen is needed for the citric acid cycle only indirectly, to regenerate the
o!idi&ed forms of :AD
;
and 0AD, which are substrates of the citric acid cycle'
13-14 (a) All the other cells can carry out o!idati"e phosphorylation to generate additional
ATP'
13-15 (c)
13-16 #hoice (d) is the correct answer' -actate is a metabolic dead end and it cannot be utili&ed
any further by cell' Pyru"ate, on the other hand, can either be con"erted to acetyl #oA
and metaboli&ed in the citric acid cycle or con"erted to glucose by a process called
gluconeogenesis' #on"ersion of lactate to pyru"ate uses up :AD
;
(making choice (b)
incorrect) and generates :AD%, but the reason animal cells make lactate in the first place
is that they are suffering from an e!cess of :AD% (and thus choice (a) is incorrect)'
#hoice (c) is incorrect since no mammal can sur"i"e without o!ygen' The amount of heat
generated by con"erting lactate to pyru"ate is negligible, so choice (e) is incorrect'
13-17 A' :AD% :AD
;
'
+' Bnder anaerobic conditions, it is the only means of regenerating the :AD
;

re(uired 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 o!ygen because less usable
energy can be har"ested from glucose in the absence of o!ygen' 6egardless of the
presence of o!ygen in the en"ironment, cells need about the same amount of energy in
the form of ATP' An the absence of o!ygen, a glucose molecule yields only $ ATP
molecules and thus many glucose molecules must be consumed to satisfy the energetic
needs of the cell' An the presence of o!ygen, a glucose molecule yields about )9 ATP
molecules'
13-19 AH$J +H2J #H4J DH)
220
13-20 #hoice (b) is the correct answer' This is another way of stating that the energetically
fa"orable o!idation of glyceraldehyde )*phosphate pro"ides sufficient energy to
ultimately dri"e the energy*re(uiring step of ATP synthesis from ADP' #hoice (a) is
untrue@ :AD% is an end product of the reaction G*)*P to 2,)*PG and therefore high (not
low) le"els of it would inhibit the reaction' #hoice (c) is untrue@ the reactions do not
in"ol"e the )*phosphate group on glyceraldehyde )*phosphate at all' #hoice (d) is untrue,
since the cysteine on the en&yme is important in making a co"alent intermediate with the
substrate and is not o!idi&ed by :AD
;
' #hoice (e) is untrue, since if the reaction had an
o"erall positi"e G, it could not be used to power the energetically unfa"orable reactions
of ATP and :AD% synthesis'
13-21 #hoice (b) is the correct answer' The phosphorylation and release of the product from the
normal en&yme is possible because the high*energy thioester bond formed between the
o!idi&ed substrate and en&yme can be attacked by a phosphate molecule' Af the bond
between o!idi&ed substrate and en&yme is a much lower*energy bond, the en&yme will
not be able to transfer the o!idi&ed substrate to a phosphate group, and substrate and
en&yme will remain co"alently bound' #hoices (a), (c), and (d) could not happen, as none
of the bonds in the substrate molecule is reacti"e enough to be broken by a phosphate
group' #hoice (e) would not happen because the en&yme does not bind ATP'
13-22 #hoice (b) is the correct answer' 8ome food molecules can enter the citric acid cycle at
points other than acetyl #oA, but only molecules that enter as acetyl #oA are completely
o!idi&ed to carbon dio!ide' Acetyl #oA is made in the mitochondrial matri!, not the
intermembrane space, so choice (a) is incorrect' #hoice (c) is incorrect since no ATP is
directly generated by the con"ersion of pyru"ate to acetyl #oAJ carbon dio!ide and
:AD% are the only other products of the reaction' #hoice (d) is incorrect as the
breakdown of fatty acids to produce acetyl #oA occurs in the mitochondrial matri!'
#hoice (e) is incorrect since acetyl #oA is not an intermediate in the anaerobic
fermentation reaction that co"erts pyru"ate to lactate'
13-23 (+) This will produce $ :AD%, $ 0AD%$, and ) acetyl #oA on complete o!idation'
+ecause of the double bond in A, this fatty acid will produce $ :AD% and )
acetyl #oA, but only 2 0AD%$, since the initial o!idation step using 0AD
;
that
reduces the two*carbon unit ?#%$#%$? to ?#%=#%? is not needed for two*
carbon units already containing a double bond' 8o although the amount of acetyl
#oA entering the citric acid cycle will be the same for A and +, fewer reducing
e(ui"alents will e"entually enter the electron transport chain from the o!idation of
A, thus less ATP will be produced'
13-24 (c)
13-25 nly small amounts of o!aloacetate are re(uired relati"e to the amount of acetyl #oA
metaboli&ed because o!aloacetate is regenerated after e"ery round of the citric acid cycle'
221
13-26 #hoice (b) is the correct answer' The citric acid cycle generates high*energy electrons
that are passed to :AD
;
to form :AD%' :AD% then donates these electrons to the
electron transport chain that dri"es o!idati"e phosphorylation, regenerating the :AD
;

needed to keep the citric acid cycle going' The electrons from :AD% are passed "ia the
electron transport chain to o!ygen'
13-27 #hoices (b) and (d) are correct' To get the cycle turning you need water, :AD
;
, GDP,
phosphate, 0AD, coen&yme A, and at least one intermediate of the citric acid cycle, which
are all pro"ided in choice (d)' An addition, choice (b) will be sufficient because
o!aloacetate reacts with acetyl #oA to release a molecule of coen&yme A, which can then
be reused' #hoice (c) would produce a small amount of #$ initially from the added
citrate, but the cycle could not continue since citrate has to go through a step re(uiring
coen&yme A to complete the cycle' #hoice (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
negati"e despite a positi"e G<' The concentration ratio of products to substrates, NAAO
N:AD%O>N.A-ON:AD
;
O, must be substantially smaller than the e(uilibrium constant C to
o"ercome a positi"e G< "alue'
13-29 The two reasons are to keep the cycle going and to har"est more energy' (2) The cycle
must regenerate o!aloacetate, which acts catalytically in the citric acid cycle to aid in the
o!idation of many acetyl groups' ($) Additional reactions are re(uired to more fully
o!idi&e the carbons and har"est energy by producing se"eral acti"ated carrier molecules,
including GTP, 0AD%$, and :AD%'
13-30 #hoice (e) is the correct answer' /lectrons passing along the electron transport chain
mo"e to successi"ely lower energy states' #hoice (a) is untrue as electrons from 0AD%$
can be used as well' #hoice (b) is untrue as the two mechanisms of coupling o!idation to
phosphorylation are (uite different' !idati"e phosphorylation in"ol"es the o!idation of
:AD% to :AD
;
by proteins of the electron transport chain' /lectron transport then
causes the formation of a proton gradient across a membrane, which dri"es ATP
synthesis' An contrast, glyceraldehyde )*phosphate dehydrogenase action in"ol"es the
reduction of :AD
;
to :AD% and uses the G of glyceraldehyde o!idation to form a
high*energy bond that can be attacked directly by a phosphate group' #hoice (c) is
untrue, as electron transport occurs in the plasma membrane of procaryotes' #hoice (d) is
untrue, as molecular o!ygen acts as an acceptor, not a donor, for electrons'
222
13-31 #hoice (e) is the correct answer' Af the inner mitochondrial membrane became permeable
to protons, the electron transport chain would continue to o!idi&e :AD% to :AD
;
,
transport electrons and pump protons, so the consumption of o!ygen would not fall
(choice (d))' %owe"er, the energy stored by the protons would be immediately dissipated
as heat when they flowed back across the membrane and thus could not dri"e the
synthesis of ATP' +ut, :AD% would not build up (choice (b)), and the citric acid cycle
and glycolysis would continue (and thus #$ would still be produced, contrary to choice
(c))' 8ince glycolysis and the citric acid cycle produce $ molecules of ATP and one
molecule of GTP (which can be con"erted to ATP), respecti"ely, ATP production would
not completely cease (choice (a)), but it would be "ery much less than normal'
13-32 !idati"e phosphorylation is a process that occurs in the inner membrane of
mitochondria' At re(uires an electron*transport chain that operates on the high*energy
electrons taken from the acti"ated 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
+
' 8ince their concentration is much hiher outside than inside
the mitochondria, the flow of protons, or H
+
, down the concentration gradient is
energetically "ery !a"orab#e and can thus be coupled to the production of ATP from ADP'
Thus, o!idati"e phosphorylation refers to the o!idation of NADH and FADH2 molecules
and the phosphorylation of AD$' Without this process, the yield of ATP from each
glucose molecule would be se"ere#% decreased'
13-33 A' /!periments showed that minced pigeon muscles contained large amounts of
pyru"ate, but o!idi&ed this compound rather slowly, so that little o!ygen was
consumed and little carbon dio!ide was produced' When a tiny amount of
o!aloacetate was added to such muscle preparations, large amounts of o!ygen and
carbon dio!ide were consumed and produced, respecti"ely' Af the added
o!aloacetate was simply being o!idi&ed fully, the o!ygen consumption and
carbon dio!ide production would be e!pected to increase only slightly, but instead
the large amount of o!ygen consumed suggested that each molecule of added
o!aloacetate aided in the o!idation of many molecules of some other substance'
+' Analogous e!periments showed that the addition of se"eral other compounds, like
succinate and fumarate, had the same conse(uences as adding o!aloacetate' This
was interpreted as e"idence that these compounds are intermediates in the same
pathway and can be con"erted into, or are deri"ed from, o!aloacetate, which was
later demonstrated more directly'
13-34 #hoice (d) is the correct answer' +oth starch and glycogen are storage polymers of
glucose' #hoice (a) is false, since both plants and animals can also store food as fats and
oils' #hoice (b) is false, as although glycogen synthesis re(uires ATP, no ATP is generated
by its hydrolysis to monomers (so the number of branch points is irrele"ant to energy
storage)' #hoice (c) is false, as animal cells cannot do this' #hoice (e) is false, as the use
of protein for energy occurs only under star"ation conditions'
223
13-35 #hoice (b) is the answer' The breakdown of glycogen to glucose does not re(uire o!ygenJ
the glucose can then enter glycolysis and generate ATP by a fermentation process that
produces lactic acid' An contrast, fats are broken down to acetyl #oA that must enter the
citric acid cycle, which re(uires o!ygen to keep turning' #hoice (a) is incorrect, as a
gram of glycogen (wet or dry) produces less energy than a gram of fat' #hoice (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' #hoice (d) is incorrect, as the
actual mass of glycogen re(uired to store the same amount of energy is si!*fold greater
than the amount of fat' #hoice (e) is incorrect, as fats can be carried in the bloodstream' Af
the energy stored in glycogen is re(uired by other cells, glycogen is broken down to
glucose, and the glucose is then released into the bloodstream'
13-36 #hoice (c) is the correct answer' An o!idati"e metabolism, fatty acids can only be
con"erted to acetyl #oA, which is completely o!idi&ed to carbon dio!ide through the
citric acid cycle' An addition, bacteria can use some of this acetyl #oA as a source of
carbon atoms to replenish the citric acid cycle, whereas animals cannot' #hoices (a), (b),
and (e) are incorrect, since glucose and fructose can be con"erted to pyru"ate, and hence
to citric acid cycle intermediates, in both animal and bacterial cells, while choice (d) is
incorrect, since carbon dio!ide cannot be used as a main carbon source by either
nonphotosynthetic bacteria or animal cells'
13-37 (c) Pyru"ate cannot be con"erted into :AD%, but it can be con"erted into the other
metabolites in one or two steps'
13-38 A carbon atom in a #$ molecule in the atmosphere e"entually becomes a part of one of
the en&ymes that cataly&es glycolysis in one of your cells' The #$ first enters a cell in a
corn leaf where photosynthesis fi!es the carbon to make it part of a sugar molecule,
which tra"els from the leaf to an ear of corn where it is stored as part of a polysaccharide
star&h molecule in the corn seed' Lou then eat a corn chip made from the corn seed'
Lou digest the corn seed, and the free #'&ose tra"els in your bloodstream, e"entually
being taken up by a li"er cell and stored as #%&oen' When re(uired, this storage
molecule breaks down into glucose*2*phosphate, which enters the glycolytic pathway'
Glycolysis produces p%r'"ate, which is con"erted into acetyl #oA, which enters the
&itri& a&i( &%&#e' 8e"eral intermediates in this process can pro"ide the carbon skeleton
for production of amino a&i(s, which are then incorporated into the en&ymes that
cataly&e steps in glycolysis'
224