Essential Cell Biology 5th edition
414 CHAPTER 12 Transport Across Cell MembranesFigure 12–35 An action potential istriggered by a depolarization of aneuron’s plasma membrane. The restingmembrane potential in this neuron is –60 mV,and a stimulus that depolarizes the plasmamembrane to about –40 mV (the thresholdpotential) is applied. This depolarizingstimulus is sufficient to open voltagegatedNa + channels in the membrane andthereby trigger an action potential. As themembrane rapidly depolarizes further, themembrane potential (red curve) swings pastzero, reaching +40 mV before it returnsto its resting negative value as the actionpotential terminates. The green curve showshow the membrane potential would simplyhave relaxed back to the resting value afterthe initial depolarizing stimulus if there hadbeen no amplification by voltage-gated ionchannels in the plasma membrane.Figure 12–36 A voltage-gated Na +channel can flip from one conformationto another, depending on the membranepotential. When the membrane is at restand highly polarized, positively chargedamino acids in the voltage sensors of thechannel (red bars) are oriented by themembrane potential in a way that keepsthe channel in its closed conformation.When the membrane is depolarized,the voltage sensors shift, changing thechannel’s conformation so the channel hasa high probability of opening. But in thedepolarized membrane, the inactivatedconformation is even more stable than theopen conformation, and so, after a briefperiod spent in the open conformation,the channel becomes temporarilyinactivated and cannot open. The redarrows indicate the sequence that followsa sudden depolarization, and the blackarrow indicates the return to the originalconformation after the membrane hasrepolarized.plasma membrane potential (mV)+400–40–600 1 2time (msec)STIMULUSACTIONPOTENTIALthresholdpotentialresting membranepotentialmembrane has shifted from its resting value of about –60 mV to about+40 mV (Figure 12–35).The voltage of +40 ECB5 mV is E12.31/12.35close to the membrane potential at which theelectrochemical driving force for movement of Na + across the membraneis zero—that is, the effects of the membrane potential and the concentrationgradient for Na + are equal and opposite; therefore Na + has no furthertendency to enter or leave the cell.If these voltage-gated channels continued to respond to the depolarizedmembrane potential, the cell would get stuck with most of its Na + channelsopen. The cell is saved from this fate because voltage-gated Na +channels have an automatic inactivating mechanism—a kind of “timer”that causes them to rapidly adopt (within a millisecond or so) a specialinactivated conformation in which the channel is closed, even thoughthe membrane is still depolarized. The Na + channels remain in this inactivatedstate until the membrane potential has returned to its resting,negative value. A schematic illustration of these three distinct states ofthe voltage-gated Na + channel—closed, open, and inactivated—is shownin Figure 12–36. How they contribute to the rise and fall of an actionpotential is shown in Figure 12–37.During an action potential, voltage-gated Na + channels do not act alone.The depolarized axonal membrane is helped to return to its restingpotential by the opening of voltage-gated K + channels. These also openin response to depolarization, but not as promptly as the Na + channels,and they stay open as long as the membrane remains depolarized. As thelocal depolarization reaches its peak, K + ions (carrying positive charge)therefore start to flow out of the cell, down their electrochemical gradient,plasmamembraneEXTRACELLULARSPACECYTOSOL– – –– – –+++ + ++ + +INACTIVATED++++CLOSED+ + ++ + +–––RECOVERY ANDMEMBRANEREPOLARIZATIONREFRACTORYPERIODvoltage sensors++–––voltage-gatedNa + channelARRIVAL OFACTION POTENTIAL– – –– – –+++ + ++ + +OPEN++membraneat restmembranedepolarized
Ion Channels and Nerve Cell Signaling415membranepotential (mV)400-40closed open inactivated closed0 1 2time (milliseconds)pulse ofelectric currentFigure 12–37 Voltage-gated Na + channelschange their conformation during anaction potential. In this example, theaction potential is triggered by a brief pulseof electric current (arrow), which partiallydepolarizes the membrane, as shown in theplot of membrane potential versus time.The course of the action potential reflectsthe opening and subsequent inactivationof voltage-gated Na + channels, as shown(top). Even if restimulated, the plasmamembrane cannot produce a second actionpotential until the Na + channels havereturned from the inactivated to the closedconformation (see Figure 12–36). Until then,the membrane is resistant, or refractory, tostimulation.through these newly opened K + channels—temporarily unhindered by thenegative membrane potential that normally restrains them in the restingcell. The rapid outflow of K + through the voltage-gated K + channelsbrings the membrane back ECB5 to its e12.36/12.37resting state much more quickly thancould be achieved by K + outflow through the K + leak channels alone.Once it begins, the self-amplifying depolarization of a small patch ofplasma membrane quickly spreads outward: the Na + flowing in throughopen Na + channels begins to depolarize the neighboring region of themembrane, which then goes through the same self-amplifying cycle. Inthis way, an action potential spreads outward as a traveling wave fromthe initial site of depolarization, eventually reaching the axon terminals(Figure 12–38).QUESTION 12–5Explain as precisely as you can, butin no more than 100 words, the ionicbasis of an action potential and howit is passed along an axon.Na + CHANNELSCLOSED INACTIVATED OPEN CLOSEDNa + Na +axon plasma membrane++–– –++++––++ + + –––+REPOLARIZED DEPOLARIZEDPROPAGATIONRESTING–+–++ +––––+–time = 0 (action potential triggered)Na + Na + Na + Na +++–– –+++++–++–+ +–––++++Na + CHANNELSCLOSED INACTIVATED OPEN CLOSED++++–– – +––––++ + +–+PROPAGATIONREPOLARIZEDDEPOLARIZEDRESTINGtime = 1 millisecond (action potential travels)Na–+ – – +Na ++––+Figure 12–38 An action potentialpropagates along the length of anaxon. The changes in the Na + channelsand the consequent flow of Na + acrossthe membrane (red arrows) alters themembrane potential and gives rise tothe traveling action potential, as shownhere and in Movie 12.12. The region ofthe axon with a depolarized membraneis shaded in blue. Note that an actionpotential can only travel forward; thatis, away from the site of depolarization.This is because Na + channel inactivationin the aftermath of an action potentialprevents the advancing front ofdepolarization from spreading backward(see also Figure 12–37).
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Ion Channels and Nerve Cell Signaling
415
membrane
potential (mV)
40
0
-40
closed open inactivated closed
0 1 2
time (milliseconds)
pulse of
electric current
Figure 12–37 Voltage-gated Na + channels
change their conformation during an
action potential. In this example, the
action potential is triggered by a brief pulse
of electric current (arrow), which partially
depolarizes the membrane, as shown in the
plot of membrane potential versus time.
The course of the action potential reflects
the opening and subsequent inactivation
of voltage-gated Na + channels, as shown
(top). Even if restimulated, the plasma
membrane cannot produce a second action
potential until the Na + channels have
returned from the inactivated to the closed
conformation (see Figure 12–36). Until then,
the membrane is resistant, or refractory, to
stimulation.
through these newly opened K + channels—temporarily unhindered by the
negative membrane potential that normally restrains them in the resting
cell. The rapid outflow of K + through the voltage-gated K + channels
brings the membrane back ECB5 to its e12.36/12.37
resting state much more quickly than
could be achieved by K + outflow through the K + leak channels alone.
Once it begins, the self-amplifying depolarization of a small patch of
plasma membrane quickly spreads outward: the Na + flowing in through
open Na + channels begins to depolarize the neighboring region of the
membrane, which then goes through the same self-amplifying cycle. In
this way, an action potential spreads outward as a traveling wave from
the initial site of depolarization, eventually reaching the axon terminals
(Figure 12–38).
QUESTION 12–5
Explain as precisely as you can, but
in no more than 100 words, the ionic
basis of an action potential and how
it is passed along an axon.
Na + CHANNELS
CLOSED INACTIVATED OPEN CLOSED
Na + Na +
axon plasma membrane
+
+
–
– –
+
+
+
+
–
–
+
+ + + –
–
–
+
REPOLARIZED DEPOLARIZED
PROPAGATION
RESTING
–
+
–
+
+ +
–
–
–
–
+
–
time = 0 (action potential triggered)
Na + Na + Na + Na +
+
+
–
– –
+
+
+
+
+
–
+
+
–
+ +
–
–
–
+
+
+
+
Na + CHANNELS
CLOSED INACTIVATED OPEN CLOSED
+
+
+
+
–
– – +
–
–
–
–
+
+ + +
–
+
PROPAGATION
REPOLARIZED
DEPOLARIZED
RESTING
time = 1 millisecond (action potential travels)
Na
–
+ – – +
Na +
+
–
–
+
Figure 12–38 An action potential
propagates along the length of an
axon. The changes in the Na + channels
and the consequent flow of Na + across
the membrane (red arrows) alters the
membrane potential and gives rise to
the traveling action potential, as shown
here and in Movie 12.12. The region of
the axon with a depolarized membrane
is shaded in blue. Note that an action
potential can only travel forward; that
is, away from the site of depolarization.
This is because Na + channel inactivation
in the aftermath of an action potential
prevents the advancing front of
depolarization from spreading backward
(see also Figure 12–37).