Essential Cell Biology 5th edition
396 CHAPTER 12 Transport Across Cell MembranesQUESTION 12–1A simple enzyme reaction can bedescribed by the equationE + S ↔ ES ↔ E + P, where E isthe enzyme, S the substrate,P the product, and ES the enzyme–substrate complex.A. Write a corresponding equationdescribing the workings of atransporter (T) that mediates thetransport of a solute (S) down itsconcentration gradient.B. What does this equation tell youabout the function of a transporter?C. Why would this equation be aninappropriate choice to representthe function of a channel?Passive Transporters Move a Solute Along ItsElectrochemical GradientAn important example of a transporter that mediates passive transportis the glucose transporter in the plasma membrane of many mammaliancell types. The protein, which consists of a polypeptide chain that crossesthe membrane at least 12 times, can adopt several conformations—andit switches reversibly and randomly between them. In one conformation,the transporter exposes binding sites for glucose to the exterior of thecell; in another, it exposes the sites to the cell interior.Because glucose is uncharged, the electrical component of its electrochemicalgradient is zero. Thus the direction in which it is transported isdetermined by its concentration gradient alone. When glucose is plentifuloutside cells, as it is after a meal, the sugar binds to the transporter’sexternally displayed binding sites; if the protein then switches conformation—spontaneouslyand at random—it will carry the bound sugarinward and release it into the cytosol, where the glucose concentrationis low (Figure 12–9). Conversely, when blood glucose levels are low—asthey are when you are hungry—the hormone glucagon stimulates livercells to produce large amounts of glucose by the breakdown of glycogen.As a result, the glucose concentration is higher inside liver cells thanoutside. This glucose can bind to the internally displayed binding siteson the transporter. When the protein then switches conformation in theopposite direction—again spontaneously and randomly—the glucose willbe transported out of the cells and made available for import by other,energy-requiring cells. The net flow of glucose can thus go either way,according to the direction of the glucose concentration gradient acrossthe plasma membrane: inward if more glucose is binding to the transporter’sexternally displayed sites, and outward if the opposite is true.Although passive transporters themselves play no part in controllingthe direction of solute transport, they are highly selective in terms ofwhich solutes they will move. For example, the binding sites in the glucosetransporter bind only D-glucose and not its mirror image L-glucose,which the cell cannot use as an energy source.Pumps Actively Transport a Solute Against ItsElectrochemical GradientCells cannot rely solely on passive transport to maintain the properbalance of solutes. The active transport of solutes against their electrochemicalgradient is essential to achieving the appropriate intracellularglucoseEXTRACELLULARSPACEcellmembraneconcentrationgradientCYTOSOLglucose transporterglucose-binding siteFigure 12–9 Conformational changes in a transporter mediate the passive transport of a solute such as glucose. The transporteris shown in three conformational states: in the outward-open state (left), the binding sites for solute are exposed on the outside; inthe inward-open state (right), the sites are exposed on the inside of the bilayer; and in the occluded state (center), the sites are notaccessible from either side. The transition between the states occurs randomly, is completely reversible, and—most importantly forthe function of the transporter shown—does not depend on whether the solute-binding site is occupied. Therefore, if the soluteconcentration is higher on the outside of the bilayer, solute will bind more often to the transporter in the outward-open conformationthan in the inward-open conformation, and there will be a net transport of glucose down its concentration gradient.ECB5 e12.09/12.09
Transporters and Their Functions397cellmembraneLIGHTFigure 12–10 Pumps carry out activetransport in three main ways. The activelytransported solute is shown in gold, and theenergy source is shown in red.electrochemicalgradientPATPGRADIENT-DRIVENPUMPADPATP-DRIVENPUMPLIGHT-DRIVENPUMPionic composition and for importing solutes that are at a lower concentrationoutside the cell than inside. For these purposes, cells depend ontransmembrane pumps, which can carry out active transport in threemain ways (Figure 12–10): (i) gradient-driven pumps link the uphilltransport of one solute ECB5 across e12.10/12.10a membrane to the downhill transport ofanother; (ii) ATP-driven pumps use the energy released by the hydrolysisof ATP to drive uphill transport; and (iii) light-driven pumps, which arefound mainly in bacterial cells, use energy derived from sunlight to driveuphill transport, as discussed in Chapter 11 for bacteriorhodopsin (seeFigure 11–28).These different forms of active transport are often linked. Thus, in theplasma membrane of an animal cell, an ATP-driven Na + pump transportsNa + out of the cell against its electrochemical gradient; this Na + can thenflow back into the cell, down its electrochemical gradient, through variousNa + gradient-driven pumps. The influx of Na + through these gradientdrivenpumps provides the energy for the active transport of many othersubstances into the cell against their electrochemical gradients. If theATP-driven Na + pump ceased operating, the Na + gradient would soon rundown, and transport through Na + gradient-driven pumps would come toa halt. For this reason, the ATP-driven Na + pump has a central role in theactive transport of small molecules across the plasma membrane of animalcells. Plant cells, fungi, and many bacteria use ATP-driven H + pumpsin an analogous way: in pumping H + out of the cell, these proteins createan electrochemical gradient of H + across the plasma membrane that issubsequently harnessed for solute transport, as we discuss later.The Na + Pump in Animal Cells Uses Energy Supplied byATP to Expel Na + and Bring in K +The ATP-driven Na + pump plays such a central part in the energy economyof animal cells that it typically accounts for 30% or more of theirtotal ATP consumption. This pump uses the energy derived from ATPhydrolysis to transport Na + out of the cell as it carries K + in. The pump istherefore sometimes called the Na + -K + ATPase or the Na + -K + pump.During the pumping process, the energy from ATP hydrolysis fuels a stepwiseseries of protein conformational changes that drives the exchangeof Na + and K + ions. As part of the process, the phosphate group removedfrom ATP gets transferred to the pump itself (Figure 12−11).The transport of Na + ions out, and K + ions in, takes place in a cycle inwhich each step depends on the one before (Figure 12−12). If any of theindividual steps is prevented from occurring, the entire cycle halts. Thetoxin ouabain, for example, inhibits the Na + pump by preventing the bindingof extracellular K + , arresting the cycle.
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Transporters and Their Functions
397
cell
membrane
LIGHT
Figure 12–10 Pumps carry out active
transport in three main ways. The actively
transported solute is shown in gold, and the
energy source is shown in red.
electrochemical
gradient
P
ATP
GRADIENT-DRIVEN
PUMP
ADP
ATP-DRIVEN
PUMP
LIGHT-DRIVEN
PUMP
ionic composition and for importing solutes that are at a lower concentration
outside the cell than inside. For these purposes, cells depend on
transmembrane pumps, which can carry out active transport in three
main ways (Figure 12–10): (i) gradient-driven pumps link the uphill
transport of one solute ECB5 across e12.10/12.10
a membrane to the downhill transport of
another; (ii) ATP-driven pumps use the energy released by the hydrolysis
of ATP to drive uphill transport; and (iii) light-driven pumps, which are
found mainly in bacterial cells, use energy derived from sunlight to drive
uphill transport, as discussed in Chapter 11 for bacteriorhodopsin (see
Figure 11–28).
These different forms of active transport are often linked. Thus, in the
plasma membrane of an animal cell, an ATP-driven Na + pump transports
Na + out of the cell against its electrochemical gradient; this Na + can then
flow back into the cell, down its electrochemical gradient, through various
Na + gradient-driven pumps. The influx of Na + through these gradientdriven
pumps provides the energy for the active transport of many other
substances into the cell against their electrochemical gradients. If the
ATP-driven Na + pump ceased operating, the Na + gradient would soon run
down, and transport through Na + gradient-driven pumps would come to
a halt. For this reason, the ATP-driven Na + pump has a central role in the
active transport of small molecules across the plasma membrane of animal
cells. Plant cells, fungi, and many bacteria use ATP-driven H + pumps
in an analogous way: in pumping H + out of the cell, these proteins create
an electrochemical gradient of H + across the plasma membrane that is
subsequently harnessed for solute transport, as we discuss later.
The Na + Pump in Animal Cells Uses Energy Supplied by
ATP to Expel Na + and Bring in K +
The ATP-driven Na + pump plays such a central part in the energy economy
of animal cells that it typically accounts for 30% or more of their
total ATP consumption. This pump uses the energy derived from ATP
hydrolysis to transport Na + out of the cell as it carries K + in. The pump is
therefore sometimes called the Na + -K + ATPase or the Na + -K + pump.
During the pumping process, the energy from ATP hydrolysis fuels a stepwise
series of protein conformational changes that drives the exchange
of Na + and K + ions. As part of the process, the phosphate group removed
from ATP gets transferred to the pump itself (Figure 12−11).
The transport of Na + ions out, and K + ions in, takes place in a cycle in
which each step depends on the one before (Figure 12−12). If any of the
individual steps is prevented from occurring, the entire cycle halts. The
toxin ouabain, for example, inhibits the Na + pump by preventing the binding
of extracellular K + , arresting the cycle.