Essential Cell Biology 5th edition
A:32 Answersin a membrane. A voltage of 150,000 V would instantlydischarge in an arc across a 1-cm-wide gap (that is, air wouldbe an insufficient insulator for this strength of field).ANSWER 12–15A. Nothing. You require ATP to drive the Na + pump.B. The ATP becomes hydrolyzed, and Na + is pumped intothe vesicles, generating a concentration gradient of Na +across the membrane. At the same time, K + is pumpedout of the vesicles, generating a concentration gradientof K + of opposite polarity. When all the K + is pumpedout of the vesicle or the ATP runs out, the pump wouldstop.C. The pump would initiate a transport cycle and thencease. Because all reaction steps must occur strictlysequentially, dephosphorylation and the accompanyingconformational switch cannot occur in the absenceof K + . The Na + pump will therefore become stuck inthe phosphorylated state, waiting indefinitely for apotassium ion. The number of sodium ions transportedwould be minuscule, because each pump moleculewould have functioned only a single time. Similarexperiments, leaving out individual ions and analyzingthe consequences, were used to determine thesequence of steps by which the Na + pump works.D. ATP would become hydrolyzed, and Na + and K + wouldbe pumped across the membrane as described in (B).However, the pump molecules that sit in the membranein the reverse orientation would be completely inactive(i.e., they would not—as one might have erroneouslyassumed—pump ions in the opposite direction)because ATP would not have access to the site onthese molecules where phosphorylation occurs, which isnormally exposed to the cytosol. ATP is highly chargedand cannot cross membranes without the help of specifictransporters.E. ATP becomes hydrolyzed, and Na + and K + are pumpedacross the membrane, as described in (B). K + , however,immediately flows back into the vesicles through theK + leak channels. K + moves down the K + concentrationgradient formed by the action of the Na + pump.With each K + that moves into the vesicle through aleak channel, a positive charge is moved across themembrane, generating a membrane potential that ispositive on the inside of the vesicles. Eventually, K +will stop flowing through the leak channels when themembrane potential balances the K + concentrationgradient. The scenario described here is a slightoversimplification: the Na + pump in mammaliancells actually moves three sodium ions out of cellsfor each two potassium ions that it pumps, therebydriving an electric current across the membrane andmaking a small additional contribution to the restingmembrane potential (which therefore corresponds onlyapproximately to a state of equilibrium for K + moving viaK + leak channels).ANSWER 12–16 Ion channels can be ligand-gated, voltagegated,or mechanically- (stress-) gated.ANSWER 12–17 The cell has a volume of 10 –12 liters(= 10 –15 m 3 ) and thus contains 6 × 10 4 calcium ions(= 6 × 10 23 molecules/mole × 100 × 10 –9 moles/liter× 10 –12 liters). Therefore, to raise the intracellular Ca 2+concentration fiftyfold, another 2,940,000 calcium ions haveto enter the cell (note that at 5 μM concentration there are3 × 10 6 ions in the cell, of which 60,000 are already presentbefore the channels are opened). Because each of the 1000channels allows 10 6 ions to pass per second, each channelhas to stay open for only 3 milliseconds.ANSWER 12–18 Animal cells drive most transport processesacross the plasma membrane with the electrochemicalgradient of Na + . ATP is needed to fuel the Na + pump tomaintain the Na + gradient.ANSWER 12–19A. If H + is pumped across the membrane into theendosomes, an electrochemical gradient of H + results,composed of both an H + concentration gradient anda membrane potential, with the interior of the vesiclepositive. Both of these components add to the energythat is stored in the gradient and that must be suppliedto generate it. The electrochemical gradient will limitthe transfer of more H + . If, however, the membrane alsocontains Cl – channels, the negatively charged Cl – in thecytosol will flow into the endosomes and diminish theirmembrane potential. It therefore becomes energeticallyless expensive to pump more H + across the membrane,and the interior of the endosomes can become moreacidic.B. No. As explained in (A), some acidification would stilloccur in their absence.ANSWER 12–20A. See Figure A12–20.B. The transport rates of compound A are proportionalto its concentration, indicating that compound A candiffuse through membranes on its own. Compound A islikely to be ethanol, because it is a small and relativelynonpolar molecule that can diffuse readily through thelipid bilayer (see Figure 12–2). In contrast, the transportrates of compound B saturate at high concentrations,indicating that compound B is transported across themembrane by some sort of membrane transport protein.Transport rates cannot increase beyond a maximal rateat which this protein can function. Compound B is likelyto be acetate, because it is a charged molecule thatcould not cross the membrane without the help of amembrane transport protein.rate of transport (µmol/min)200100compound Bcompound A00 2 4 6 8 10concentration of solute (mM)Figure A12–20
Answers A:33ANSWER 12–21 The membrane potential and the highextracellular Na + concentration provide a large inwardelectrochemical driving force and a large reservoir of Na +ions, so that mostly Na + ions enter the cell as acetylcholinereceptors open. Ca 2+ ions will also enter the cell, buttheir influx is much more limited because of their lowerextracellular concentration. (Most of the Ca 2+ that entersthe cytosol to stimulate muscle contraction is released fromintracellular stores, as we discuss in Chapter 17). Becauseof the high intracellular K + concentration and the opposingdirection of the membrane potential, there will be little ifany movement of K + ions upon opening of a cation channel.ANSWER 12–22 The diversity of neurotransmitter-gated ionchannels raises the hope of developing new drugs specificfor each channel type. Each of the diverse subtypes ofthese channels is expressed in a narrow subset of neurons.This narrow range of expression should make it possible, inprinciple, to discover or design drugs that affect particularreceptor subtypes present in a selected set of neurons, thustargeting particular brain functions with greater specificity.Chapter 13ANSWER 13–1 To keep glycolysis going, cells needto regenerate NAD + from NADH. In the absence ofoxygen, there is no efficient way to do this withoutfermentation. Without regenerated NAD + , step 6 ofglycolysis [the oxidation of glyceraldehyde 3-phosphate to1,3-bisphosphoglycerate (Panel 13–1, pp. 436–437)] couldnot occur, and the product glyceraldehyde 3-phosphatewould accumulate. The same thing would happen in cellsunable to make either lactate or ethanol: neither wouldbe able to regenerate NAD + , and so glycolysis would beblocked at the same step.ANSWER 13–2 Arsenate instead of phosphate becomesattached in step 6 of glycolysis to form 1-arseno-3-phosphoglycerate (Figure A13–2). Because of its sensitivityto hydrolysis in water, the high-energy bond is destroyedbefore the molecule that contains it can diffuse toreach the next enzyme. The product of the hydrolysis,3-phosphoglycerate, is the same product normally formedin step 7 by the action of phosphoglycerate kinase. Butbecause hydrolysis occurs nonenzymatically, the energyliberated by breaking the high-energy bond cannot becaptured to generate ATP. In Figure 13–7, therefore, thereaction corresponding to the downward-pointing arrowin step 7 would still occur, but the wheel that providesthe coupling to ATP synthesis is missing. Arsenate wastesmetabolic energy by uncoupling many phosphotransferreactions by the same mechanism, which is why it is sopoisonous.ANSWER 13–3 The oxidation of fatty acids breaks thecarbon chain down into two-carbon units at a time (acetylgroups that had become attached to CoA). Conversely,during biogenesis, fatty acids are constructed by linkingtogether acetyl groups. Most fatty acids therefore have aneven number of carbon atoms.ANSWER 13–4 Because the function of the citric acid cycleis to harvest the energy released during the oxidation, itis advantageous to break the overall reaction into as manysteps as possible (see Figure 13–1). Using a two-carboncompound (acetyl CoA), the available chemistry would bemuch more limited, and it would be impossible to generateas many intermediates.ANSWER 13–5 It is true that oxygen atoms are returnedto the atmosphere as part of CO 2 during the oxidativedegradation of glucose (see Figure 13−3). The CO 2released from the cells, however, does not contain thespecific oxygen atoms consumed as part of the oxidativephosphorylation process; these are converted into water.One can show this directly by incubating living cells in anatmosphere that includes molecular oxygen containing the18 O isotope of oxygen instead of the naturally abundantisotope, 16 O. In such an experiment, one finds that all theCO 2 released from cells contains only 16 O. Therefore, theoxygen atoms in the released CO 2 molecules do not comedirectly from the atmosphere but from organic moleculesthat the cell has first made and then oxidized as fuel (seetop of first page of Panel 13–2, pp. 442–443).ANSWER 13–6 The cycle continues because intermediatesare replenished as necessary by reactions leading intothe citric acid cycle (instead of away from it). One of themost important reactions of this kind is the conversionof pyruvate to oxaloacetate by the enzyme pyruvatecarboxylase:pyruvate + CO 2 + ATP + H 2 O → oxaloacetate + ADP + P i + 2H +This reaction feeds oxaloacetate into the citric acid cycle.It is one of the many examples of how metabolic pathwaysare carefully coordinated to work together to maintainappropriate concentrations of all metabolites required bythe cell (see Figure A13–6).ADP+ PATPCO 2pyruvatecarboxylaseoxaloacetatepyruvateacetyl CoACITRICACIDCYCLEcitrateO OCH C OHOAsO OHO –O – CH C OH + AsO3– 4 +CH 2 O PCH 2 O PH 2 OFigure A13–2H +Figure A13–6ANSWER 13–7 The carbon atoms in sugar molecules arealready partially oxidized. In contrast, only the very firstcarbon atom in the acyl chains of fatty acids is oxidized.Thus, two carbon atoms from glucose are lost as CO 2 duringthe conversion of pyruvate to acetyl CoA (see Figure 13−3),leaving only four carbons to enter the citric acid cycle,ECB5 eA13.02/A13.02ECB5 eA13.06/A13.06
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Answers A:33
ANSWER 12–21 The membrane potential and the high
extracellular Na + concentration provide a large inward
electrochemical driving force and a large reservoir of Na +
ions, so that mostly Na + ions enter the cell as acetylcholine
receptors open. Ca 2+ ions will also enter the cell, but
their influx is much more limited because of their lower
extracellular concentration. (Most of the Ca 2+ that enters
the cytosol to stimulate muscle contraction is released from
intracellular stores, as we discuss in Chapter 17). Because
of the high intracellular K + concentration and the opposing
direction of the membrane potential, there will be little if
any movement of K + ions upon opening of a cation channel.
ANSWER 12–22 The diversity of neurotransmitter-gated ion
channels raises the hope of developing new drugs specific
for each channel type. Each of the diverse subtypes of
these channels is expressed in a narrow subset of neurons.
This narrow range of expression should make it possible, in
principle, to discover or design drugs that affect particular
receptor subtypes present in a selected set of neurons, thus
targeting particular brain functions with greater specificity.
Chapter 13
ANSWER 13–1 To keep glycolysis going, cells need
to regenerate NAD + from NADH. In the absence of
oxygen, there is no efficient way to do this without
fermentation. Without regenerated NAD + , step 6 of
glycolysis [the oxidation of glyceraldehyde 3-phosphate to
1,3-bisphosphoglycerate (Panel 13–1, pp. 436–437)] could
not occur, and the product glyceraldehyde 3-phosphate
would accumulate. The same thing would happen in cells
unable to make either lactate or ethanol: neither would
be able to regenerate NAD + , and so glycolysis would be
blocked at the same step.
ANSWER 13–2 Arsenate instead of phosphate becomes
attached in step 6 of glycolysis to form 1-arseno-3-
phosphoglycerate (Figure A13–2). Because of its sensitivity
to hydrolysis in water, the high-energy bond is destroyed
before the molecule that contains it can diffuse to
reach the next enzyme. The product of the hydrolysis,
3-phosphoglycerate, is the same product normally formed
in step 7 by the action of phosphoglycerate kinase. But
because hydrolysis occurs nonenzymatically, the energy
liberated by breaking the high-energy bond cannot be
captured to generate ATP. In Figure 13–7, therefore, the
reaction corresponding to the downward-pointing arrow
in step 7 would still occur, but the wheel that provides
the coupling to ATP synthesis is missing. Arsenate wastes
metabolic energy by uncoupling many phosphotransfer
reactions by the same mechanism, which is why it is so
poisonous.
ANSWER 13–3 The oxidation of fatty acids breaks the
carbon chain down into two-carbon units at a time (acetyl
groups that had become attached to CoA). Conversely,
during biogenesis, fatty acids are constructed by linking
together acetyl groups. Most fatty acids therefore have an
even number of carbon atoms.
ANSWER 13–4 Because the function of the citric acid cycle
is to harvest the energy released during the oxidation, it
is advantageous to break the overall reaction into as many
steps as possible (see Figure 13–1). Using a two-carbon
compound (acetyl CoA), the available chemistry would be
much more limited, and it would be impossible to generate
as many intermediates.
ANSWER 13–5 It is true that oxygen atoms are returned
to the atmosphere as part of CO 2 during the oxidative
degradation of glucose (see Figure 13−3). The CO 2
released from the cells, however, does not contain the
specific oxygen atoms consumed as part of the oxidative
phosphorylation process; these are converted into water.
One can show this directly by incubating living cells in an
atmosphere that includes molecular oxygen containing the
18 O isotope of oxygen instead of the naturally abundant
isotope, 16 O. In such an experiment, one finds that all the
CO 2 released from cells contains only 16 O. Therefore, the
oxygen atoms in the released CO 2 molecules do not come
directly from the atmosphere but from organic molecules
that the cell has first made and then oxidized as fuel (see
top of first page of Panel 13–2, pp. 442–443).
ANSWER 13–6 The cycle continues because intermediates
are replenished as necessary by reactions leading into
the citric acid cycle (instead of away from it). One of the
most important reactions of this kind is the conversion
of pyruvate to oxaloacetate by the enzyme pyruvate
carboxylase:
pyruvate + CO 2 + ATP + H 2 O → oxaloacetate + ADP + P i + 2H +
This reaction feeds oxaloacetate into the citric acid cycle.
It is one of the many examples of how metabolic pathways
are carefully coordinated to work together to maintain
appropriate concentrations of all metabolites required by
the cell (see Figure A13–6).
ADP
+ P
ATP
CO 2
pyruvate
carboxylase
oxaloacetate
pyruvate
acetyl CoA
CITRIC
ACID
CYCLE
citrate
O O
C
H C OH
O
As
O OH
O –
O – C
H C OH + AsO
3– 4 +
CH 2 O P
CH 2 O P
H 2 O
Figure A13–2
H +
Figure A13–6
ANSWER 13–7 The carbon atoms in sugar molecules are
already partially oxidized. In contrast, only the very first
carbon atom in the acyl chains of fatty acids is oxidized.
Thus, two carbon atoms from glucose are lost as CO 2 during
the conversion of pyruvate to acetyl CoA (see Figure 13−3),
leaving only four carbons to enter the citric acid cycle,
ECB5 eA13.02/A13.02
ECB5 eA13.06/A13.06