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11
Transmembrane Transport of Ions
and Small Molecules

Sections 11.1

1. The partition coefficient K, the equilibrium constant for partitioning between oil and water, for
butyric acid is about 10 – 2 , and for 1,4-butanediol it is about 10 – 4. You add liposomes containing
only water to a solution with an initial concentration of 1 mM butyric acid and 100 mM 1,4-
butanediol outside the liposomes. What is the relative rate of diffusion of the two substances
into the liposome interior?
a. Butyric acid diffuses into the liposome 100-fold faster than butanediol.
b. The two diffuse into the liposome at equal rates.
c. Butanediol diffuses into the liposome 100-fold faster than butanediol.
d. Not enough information is provided to permit an answer.

Ans꞉ b
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2. Classes of membrane transport proteins include all of the following EXCEPT꞉


a. ATP-powered pumps.
b. ion channels.
c. protein translocons.
d. transporters.

Ans꞉ c
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3. The glucose uniporter GLUT1 has a Km of 1.5 mM for D-glucose and 30 mM for D-galactose.
At a concentration of 5 mM for each, what is the rate of glucose transport relative to galactose
transport? The Vmax may be assumed to be the same for both.
a. 3.7-fold slower
b. equal
c. 5.5-fold faster
d. 20-fold faster

Ans꞉ c
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4. Both ethanol and glycine are small molecules of approximately equal molecular weight.
However, the membrane is much more permeable to ethanol than glycine. What accounts for
the large difference in membrane permeability between ethanol and glycine?
Ans꞉ Ethanol is a small alcohol; glycine is a small amino acid. Glycine, like all amino acids, is a
zwitterion. At neutral pH, the amino group of glycine is positively charged and the carboxyl
group is negatively charged. Charged groups are impermeable to lipid bilayers.
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5. In a laboratory activity, red-labeled glucose, blue-labeled water, and green-labeled ethanol


are added to a solution placed over an artificial, pure phospholipid membrane. Which colors will
be observed on the other side of the membrane after 10 minutes?
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a. red
b. blue
c. green
d. b and c

Ans꞉ d
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6. In which of the following cases is energy NOT needed for transmembrane transport?
a. Lysine moves into the cell against its concentration gradient via the Na+/lysine symporter.
b. Potassium ions (K+) move out of the cell down the K+ concentration gradient via potassium
channels.
c. Glucose moves into the cell down its concentration gradient via a glucose uniporter.
d. The second and third answers are correct.
e. all of the above

Ans꞉ d
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Section 11.2

7. How does uniport transport compare with simple diffusion?


a. Similar to simple diffusion, uniport transport is nonspecific.
b. Uniport transport is slower but more specific than simple diffusion.
c. Uniport transport is much faster and more specific than simple diffusion.
d. Simple diffusion is reversible but uniport transport is not.

Ans꞉ c
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8. Aquaporins are꞉
a. -barrel proteins in the outer membrane of bacteria.
b. ABC proteins.
c. symporters.
d. water channels.

Ans꞉ d
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9. When computing the osmotic pressure that must be placed across the membrane to stop the
flow of water, what is the glucose osmotic equivalent of 1 M CaCl2?
a. 1 M
b. 2 M
c. 3 M
d. 4 M

Ans꞉ c
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10. Describe how aquaporins facilitate the movement of water across the plasma membrane.

Ans꞉ Aquaporins are water-channel proteins that specifically increase the permeability of
biomembranes to water. The level of aquaporin 2 is rate-limiting for water transport by the
kidney and is essential for resorption of water in the kidney.
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11. What is the approach that plants use to respond to differences in osmotic pressure between
the inside and outside of the cell?
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Ans꞉ Plants have rigid cell walls. The normal concentration of solutes is higher inside the plant
vacuole than in the cytosol. Likewise, the solute concentration is higher in the cytosol than in
the extracellular space. Hence, the plant cell exerts pressure against the cell wall and is retained
within the cell wall.
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12. Amino acid entry into cells can occur via uniporters or symporters. If the rate of leucine
entry into the cell increases when the pH decreases, this suggests꞉
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a. leucine is being transported by a uniporter.


b. leucine is being actively pumped across the membrane against its concentration gradient.
c. leucine is being transported by a H+ symporter.
d. leucine is crossing the membrane via simple diffusion.

Ans꞉ c
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13. Glucose enters erythrocytes via a GLUT-1 uniporter. As the levels of glucose in the
bloodstream decrease between meals, what happens to the glucose in the cells?
a. Glucose leaves the cell through the GLUT-1 uniporter, traveling down the new concentration
gradient.
b. Glucose remains in the cell because uniporters can only transport in one direction.
c. Glucose remains in the cell because the GLUT-1 uniporters are gated and the gates close at
low glucose concentrations.
d. Glucose remains in the cell because it has been phosphorylated and no longer has affinity for
the GLUT-1 uniporter.

Ans꞉ d
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14. Which of the following is true about glucose transporters?


a. GLUT-2 and GLUT-4 are expressed on all cell types.
b. All the members of the GLUT family have the same affinity for glucose.
c. Cells that express GLUT-1 and GLUT-3 display more glucose uptake than cells expressing
GLUT-2 at physiological blood concentrations (5 mM).
d. GLUT proteins are specific for glucose molecules and will not transport other sugars.

Ans꞉ c
Question Type꞉ Multiple Choice
Chapter꞉ 11
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15. The osmotic potential of a hypotonic solution should cause an animal cell to _____, but with
frog oocytes prevent this effect by _____.
a. burst; not expressing aquaporins
b. shrink; not expressing aquaporins
c. burst; expressing more aquaporins
d. shrink; expressing more aquaporins

Ans꞉ a
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Section 11.3

16. Which of the four classes of ATP-powered pumps share overall similarity꞉ several subunits,
the same general organization, and a similar function as H+ transporters?
a. ABC superfamily and P-class pumps
b. ABC superfamily and P-class pumps
c. F-class pumps and P-class pumps
d. F-class pumps and V-class pumps

Ans꞉ d
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17. The major ATP-powered pump responsible for maintaining ion gradients across the plasma
membrane of mammalian cells is꞉
a. the calmodulin-activated plasma membrane Ca2+ ATPase.
b. the sarcoplasmic reticulum Ca2+ ATPase.
c. the vacuolar F-class proton pump.
d. the plasma-membrane Na+/K+ ATPase.
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Ans꞉ d
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18. ABC superfamily proteins are thought to act as ATP-dependent flippases in transporting꞉
a. lipophilic drugs out of mammalian cells.
b. Ca2+ out of mammalian cells.
c. H+ out of mammalian cells.
d. Na+ out of mammalian cells, K+ into mammalian cells.

Ans꞉ a
Question Type꞉ Multiple Choice
Chapter꞉ 11
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19. Which statement describes the mode of action of the ABCB1 transporter (the first eukaryotic
ABC transporter to be recognized)?
a. During transport, the ligand binding site is alternately exposed to the exoplasmic and the
cytoplasmic side of the membrane.
b. During transport, a conserved aspartate residue is phosphorylated.
c. This class of pumps transports only H+ ions.
d. This transporter acts as a chloride channel.

Ans꞉ a
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20. Which of the following statement(s) is (are) true of V-class proton pumps?
a. They are ATPases.
b. They are present in membranes of plant vacuoles.
c. They serve to decrease the pH inside a lysosome.
d. all of the above
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Ans꞉ d
Question Type꞉ Multiple Choice
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21. Describe in general terms how the muscle Ca2+ ATPase pumps Ca2+ ions from the cytosol into
the sarcoplasmic reticulum (SR).

Ans꞉ A P-class Ca2+ ATPase located in the SR membrane of the skeletal muscle pumps Ca2+ from
the cytosol into the lumen of the SR. This pump is the major integral membrane protein in SR
membranes and hence can be readily purified. Ca2+ pumping from the cytosol to the SR involves
a series of ordered steps and two conformational states of the pump, termed E1 and E2. In the
E1 state, the pump binds two cytosolic Ca2+ ions and ATP. The ATP is cleaved to ADP with
phosphorylation of an aspartic acid with a high-energy acyl phosphate. Next, a reduction in the
energy state of the aspartate acyl phosphate produces a conformational change in the protein
from the E1 to E2 state. In the E2 state, the affinity of the pump for Ca2+ is 1000-fold less and
both calcium ions are released into the SR lumen. With dephosphorylation of the pump, there is
a second conformational change and the pump reverts again to the E1 state.
Question Type꞉ Essay
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22. What evidence suggests that all of the P-class ion pumps evolved from a common ancestor
even though they now transport different ions?

Ans꞉ The catalytic subunits of all P-class ion pumps share a similar sequence, including a
conserved aspartate residue that is phosphorylated during transport. The 3-D structures of the
catalytic subunits are similar for those P-class ion pump structures that are known.
Question Type꞉ Essay
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23. What is the basic structural organization of an ABC superfamily transport protein?
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Ans꞉ All members of the ABC superfamily of proteins contain two transmembrane domains and
two cytosolic ATP-binding domains, which couple ATP-hydrolysis to solute movement. The
transmembrane domains associate with each other. A transmembrane domain and associated
cytosolic domain together form what may be thought of as a monomer. In nature, these two
domains may be present in one polypeptide and the overall ABC transport protein then consists
of two polypeptides. Alternatively, the four domains may be present as separate subunits or in
some cases fused as a single protein.
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24. Explain why ATP-powered proton pumps cannot by themselves acidify the lumen of the
lysosome.

Ans꞉ The V-class proton pump responsible for acidifying the lumen of the lysosome, where the
net movement of electric charge occurs during transport, is electrogenic. As each proton is
pumped into the lumen, it leaves behind a negatively charged ion in the cytosol. Positive and
negative ions attract each other across the membrane of the lysosome, which causes an electric
potential. Soon, pumping leads to a buildup of protons in the lumen, which repels other protons.
At this point, a significant transmembrane proton concentration gradient cannot be established.
Thus, to generate an acidic environment necessary to breakdown components imported into the
lysosome, proton transport must be accompanied by an equal number of anions, in this case in
the form of Cl– , into the lumen.
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25. The many pumps present in the membrane have interconnected and important functions.
Which of the following functions is NOT correct?
a. A P-class pump is involved in muscle relaxation.
b. A V-class pump is involved in maintaining the low pH of lysosomes.
c. A P-class pump is involved in keeping the cytosolic levels of calcium low.
d. A P-class pump is involved in keeping the cytosolic levels of potassium low.

Ans꞉ d
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26. Which of the following is true about cystic fibrosis?


a. The normal CFTR channel protein is a chloride uniporter, but CF patients have a mutation
that makes it require ATP.
b. CFTR mutations result in a mutated protein, which pumps too many chloride ions out of the
cell.
c. There are no treatments for patients with CF that are specifically targeted to the CFTR
receptor.
d. Patients with the most common mutation make a CFTR protein that is not present in the
membrane.

Ans꞉ d
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Sections 11.4

27. The resting membrane potential in animal cells depends largely on nongated _____ channels.
a. Ca2+
b. H+
c. K+
d. Na+

Ans꞉ c
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28. The magnitude of the membrane electrical potential is calculated by꞉


a. the Nernst equation.
b. the Michaelis–Menten equation.
c. the Faraday equation.
d. the Bose–Einstein equation.

Ans꞉ a
Question Type꞉ Multiple Choice
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29. Ion channels achieve selectivity in transport through all of the following mechanisms
EXCEPT꞉
a. evolution from distinct and different parent proteins.
b. divergent evolution from a single type of channel protein.
c. low activation energy for selective passage of the dehydrated ion.
d. use of P segments to form an ion selectivity filter.

Ans꞉ a
Question Type꞉ Multiple Choice
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30. Why are nongated channels important in the generation of a negative electric potential
(voltage) of 50–70 mV inside the cell with respect to the outside?

Ans꞉ If there is no ion movement across the membrane, there is no membrane potential. This is
true even if there is a difference in ion concentrations on either side of the membrane. The
presence of K+ channels that are usually open allows for ion movement from the inside of the
cell to the outside and the creation of a negative membrane potential inside the cell.
Question Type꞉ Essay
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31. Calculate the G for the movement of Na+ from inside a typical mammalian cell to outside.

Ans꞉ As summarized in Table 11-2 on text page 448, the cellular concentration of Na+ is 12 mM
and the blood concentration is 145 mM. The inside of the cell has a negative electric potential of
–70 mV. Therefore, for K+ the G for Na+ export will have a positive value for the Gc term (Na+
is being moved against a concentration gradient) and a positive value for the Gm term (Na+ is
being moved against the negative electric potential inside the cell). The sum of these two terms
is +3.06 kcal/mol. The Na+/K+ ATPase uses energy from ATP cleavage to export 3 Na+ for every 2
K+ imported.
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32. Describe the patch-clamping technique. Why do investigators often use frog oocytes in their
patch-clamping investigations?

Ans꞉ By pressing a special electrode against a “patch” of plasma membrane and forming a tight
seal, an investigator can “clamp” the voltage (or current) at a constant value and study the
opening, closing, regulation, and ion conductance of a single ion channel. The technique can be
used on whole cells or isolated membrane patches and there are advantages to using one over
the other. Frog oocytes are often used in patch-clamping experiments because they are readily
available and large enough to microinject in vitro transcribed mRNAs encoding channel proteins,
which will be expressed on the cell surface. Since frog oocytes normally do not express any
channel proteins of their own, only the channel protein expressed from the microinjected mRNA
will be present, allowing it to be studied in isolation.
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33. A mutant channel protein is expressed in an oocyte. Compared to patch clamping


experiments in oocytes expressing the normal channel, the length of downward deviations is
diminished by half. This indicates꞉
a. the mutant channel doesn’t make a functional channel.
b. the mutant channel doesn’t stay open as long.
c. the mutant channel opens more frequently.
d. the mutant channel behaves the same as the wild-type channel.

Ans꞉ b
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Section 11.5

34. The G calculation for the two-Na+/one-glucose symporter includes which of the following
terms?
a. RT ln [glucosein ]/[glucoseout]
b. 2RT ln [Na+in ]/[Na+out]
c. 2FE
d. all of the above

Ans꞉ d
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35. How does inhibition of the Na+/K+ ATPase increase the force of heart muscle contraction?
a. It increases cytosolic Na+ and therefore decreases Ca2+ export.
b. It increases cytosolic K+ and therefore decreases Ca2+ export.
c. It decreases cytosolic Na+ and therefore decreases Ca2+ export.
d. It decreases cytosolic K+ and therefore decreases Ca2+ export.

Ans꞉ a
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36. What is the expected effect on salt accumulation in the plant vacuole of a mutation in the
plant vacuolar ATPase that decreases the H+ concentration in the vacuole?
a. increased Na+ accumulation
b. decreased Na+ accumulation
c. no effect
d. complete seed germination failure

Ans꞉ b
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37. Propose a rationale for why the import of amino acids or sugars into cells is typically coupled
to Na+ ion import.

Ans꞉ The Na+/K+ ATPase establishes and maintains large differences in Na+ and K+ distributions
across the plasma membrane. The concentration of Na+ is low inside cells and high outside.
The membrane potential is negative. The import of Na+ is both concentration and membrane-
potential favorable and can therefore drive the import of amino acids or sugars. The exact
opposite is true for K+. Coupling to Na+ movement is accomplished by symporters.
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38. Propose a rationale for why the import of sucrose into the plant vacuole is coupled to the
export of H+ ion into the plant cytosol.

Ans꞉ The plant vacuole is acidified by proton pumps. Moreover, the proton pumps generate a
positive membrane potential in the vacuole relative to the cytosol. The export of H+ into the
cytosol is both concentration and membrane-potential favorable and can therefore drive the
import of sugars (i.e., sucrose) into the vacuole. Coupling of sucrose import to H+ export is
accomplished by antiporters.
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38. The movement of dietary glucose and sodium from the intestine into the bloodstream relies
on which of the following transport mechanisms?
a. sodium glucose symporter
b. transcellular transport
c. sodium/potassium ATPase
d. all of the above

Ans꞉ d
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Section 11.6

39. Transepithelial glucose transport uses a symporter to transport glucose up a concentration


gradient by꞉
a. coupling glucose transport to proton movement.
b. coupling glucose transport to Na+ movement.
c. coupling glucose transport to Ca2+ movement.
d. coupling glucose transport to Cl– movement.

Ans꞉ b
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40. Parietal cells acidify the stomach contents while maintaining a neutral cytosolic pH by꞉
a. exporting “excess” cytosolic OH– as HCO3– .
b. exchanging HCO3– for Cl– .
c. preserving electroneutrality by accompanying the movement of each Cl– ion into the stomach
lumen by a K+.
d. all of the above.
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Ans꞉ d
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41. Transepithelial transport requires a _____ cell layer.


a. polarized
b. sealed
c. polarized and sealed
d. permeable

Ans꞉ c
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42. How can a uniporter, GLUT2, be sufficient for entry of glucose from intestinal epithelial cells
into the bloodstream?

Ans꞉ Glucose is actually present in a higher concentration inside the intestinal epithelial cell than
in the bloodstream. Hence, a uniporter on the basolateral surface of the intestinal epithelial cell
can be effective in facilitating the entry of glucose into the bloodstream. The two-Na+/one-
glucose symporter located on the apical surface, facing the intestinal lumen, is an example of
secondary active transport generating the high glucose concentration inside the epithelial cell.
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43. The H+/K+ ATPase on the apical surface of parietal cells exports H+ and imports K+. How is
the buildup of excess K+ ions in the parietal cell cytosol prevented?

Ans꞉ A K+ channel on the apical surface of the parietal cell removes the excess K+. The outcome
of transepithelial transport is the summation of processes mediated by a number of transport
proteins.
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Chapter꞉ 11
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