CHAPTER 12

MEMBRANE TRANSPORT
2004 Garland Science Publishing
Princiles !" Me#brane Trans!r$
12-1 The most abundant intracellular cation is
(a) Na
+
.
(b) Ca
2+
.
(c) H
+
.
(d) K
+
.
(e) positively charged macromolecules.
12-2 Circle the molecule in each pair that is more likely to diuse through the lipid bilayer.
!. amino acids ben"ene
#. Cl
$
ethanol
C. glucose %N!
&. H2' '2
(. adenosine !T)
12-3 ! hungry yeast cell lands in a vat o grape *uice and begins to east on the sugars there+
producing carbon dio,ide and ethanol in the process-
C.H/2'. + 2!&) + 2)i

+ H
+
2C'2

+ 2CH0CH2'H + 2!T) + 2H2'
1nortunately+ the grape *uice is contaminated 2ith proteases (en"ymes that attack some
o the transport proteins in the yeast cell membrane)+ and the yeast cell dies. 3hich o the
ollo2ing could account or the yeast cell4s demise5
(a) To,ic buildup o carbon dio,ide inside the cell
(b) To,ic buildup o ethanol inside the cell
(c) &iusion o !T) out o the cell
(d) 6nability to import sugar into the cell
(e) 6nability to take 2ater into the cell
1%&
12-4 7or each o the ollo2ing sentences+ ill in the blanks 2ith the best 2ord or phrase
selected rom the list belo2. Not all 2ords or phrases 2ill be used8 each 2ord or phrase
should be used only once.
! molecule moves do2n its concentration gradient by
999999999999999999 transport+ but re:uires 999999999999999999
transport to move up its concentration gradient. Carrier proteins and ion
channels unction in membrane transport by providing a
999999999999999999 path2ay through the membrane or speciic polar
solutes or inorganic ions. 999999999999999999 are highly selective in
the solutes they transport+ binding the solute at a speciic site and changing
its conormation in order to transport the solute across the membrane. 'n
the other hand+ 999999999999999999 discriminate among solutes mainly
on the basis o si"e and electrical charge.
active carrier proteins ion channels
amino acid hydrophilic noncovalent
amphipathic hydrophobic passive
Carrier Pr!$eins and Their 'unc$i!ns
12-5 ;our riend isolates a ne2 species o yeast that can gro2 using either ethanol
(CH0CH2'H) or acetate (CH0C''
$
) as the yeast4s sole carbon source. He measures the
rate o uptake o these carbon sources by the yeast as a unction o concentration o
carbon source and obtains the ollo2ing graphs-
7igure </2=>
!. 6s molecule ! or # more likely to utili"e a carrier protein to mediate its transport5
3hy5
#. ?iven 2hat you kno2 about membrane transport+ is molecule ! more likely to be
ethanol or acetate5 3hy5
12-6 Name the three main types o active transport.
1&0
12-7 3hich o the ollo2ing descriptions are T%1( o the bacterial protein bacteriorhodopsin5
6t is a
(a) light=driven pump.
(b) proton pump.
(c) passive transporter.
(d) coupled transporter.
(e) transmembrane protein.
12-8 The !eroschmidt 2eed contains an !T)=driven ion pump in its vacuolar membrane that
pumps potentially to,ic heavy metal ions such as @n
2+
and )b
2+
into the vacuole. The
pump protein e,ists in a phosphorylated and an unphosphorylated orm and 2orks in a
similar 2ay to the Na
+
=K
+
pump o animal cells. To study its action+ you incorporate the
unphosphorylated orm o the protein into phospholipid vesicles containing K
+
in their
interiors. (;ou ensure that all o the protein molecules are in the same orientation in the
lipid bilayer.) 3hen you add @n
2+
and !T) to the solution outside such vesicles+ you ind
that @n
2+
is pumped into the vesicle lumen. ;ou then e,pose vesicles containing the pump
protein to the solutes as sho2n in Table /2=A!.
Table /2=A!
! # C & ( 7
'utside @n
2+
+ !T) @n
2+
@n
2+
+ !T) @n
2+
!T) !T)
6nside K
+
K
+
K
+
;ou then determine the amount o phosphorylated and unphosphorylated !T)=driven ion
pump protein in each sample. ;our results are summari"ed in Table /2=A#+ 2here B$C
indicates an absence o a type o protein and B+C indicates the presence o a type o
protein.
Table /2=A#
! # C & ( 7
phosphorylated + $ $ $ $ ++
protein present
unphosphorylated ++ ++ ++ ++ ++ $
protein present
3hat 2ould you e,pect to happen i you treat vesicles as in lane 7+ but beore
determining the phosphorylation state o the protein+ you 2ash a2ay the outside buer
and replace it 2ith a buer containing only @n
2+
5
(a) Nothing 2ill happen. (No @n
2+
2ill move into the vesicle8 no K
+
2ill move out o
the vesicle8 the phosphorylation state o the protein 2ill not change.)
(b) No @n
2+
2ill move into the vesicle8 no K
+
2ill move out o the vesicle8 the protein
2ill become unphosphorylated.
(c) ! small amount o @n
2+
2ill move into the vesicle8 no K
+
2ill move out o the
vesicle8 the phosphorylation state o the protein 2ill not change.
(d) ! small amount o @n
2+
2ill move into the vesicle8 no K
+
2ill move out o the
vesicle8 the protein 2ill become unphosphorylated.
(e) ! small amount o @n
2+
2ill move into the vesicle8 a small amount o K
+
2ill
move out o the vesicle8 the phosphorylation state o the protein 2ill not change.
1&1
12-9 (,plain 2hy Na
+
is commonly used to drive the coupled in2ard transport o nutrients in
animal cells.
12-10 'uabain inhibits the active uptake o glucose into epithelial cells by
(a) binding to the glucose=Na
+
symport.
(b) opening K
+
channels.
(c) changing the pH o the cell.
(d) increasing the intracellular concentration o Na
+
.
(e) depleting the cell o !T).
12-11 3hich o the ollo2ing statements is T%1(5
(a) !moebae have carrier proteins that actively pump 2ater molecules rom the
cytoplasm to the cell e,terior.
(b) #acteria and animal cells rely on the Na
+
$K
+
pump in the plasma membrane to
prevent lysis due to osmotic imbalances.
(c) The Na
+
$K
+
pump allo2s animal cells to thrive under conditions o very lo2 ionic
strength.
(d) The cell 2all surrounding plant cells prevents osmosis.
(e) The Na
+
$K
+
pump helps to keep both Na
+
and Cl
$
ions out o the cell.
12-12 The Ca
2+
pumps in the plasma membrane and endoplasmic reticulum are e,amples o
(a) !T)=driven pumps.
(b) coupled transporters.
(c) passive carrier proteins.
(d) ion channels.
(e) symports.
12-13 Ca
2+
pumps in the plasma membrane and endoplasmic reticulum are important or
(a) maintaining osmotic balance.
(b) preventing Ca
2+
rom altering the behavior o molecules in the cytosol.
(c) providing en"ymes in the endoplasmic reticulum 2ith Ca
2+
ions that are necessary
or their catalytic activity.
(d) maintaining a negative membrane potential.
(e) helping cells import K
+
.
12-14 &o you agree o disagree 2ith the ollo2ing statement5 (,plain your ans2er.
! symporter 2ould unction as an antiporter i its orientation in the
membrane 2ere reversed (that is+ i the portion o the protein normally
e,posed to the cytosol aced the outside o the cell instead).
1&2
12-15 ;ou have prepared lipid vesicles (spherical lipid bilayers) that contain Na
+
$K
+
pumps as
the sole membrane protein. !ll o the Na
+
$K
+
pumps are oriented in such a 2ay that the
portion o the molecule that normally aces the cytosol is in the inside o the vesicle and
the portion o the molecule that normally aces the e,tracellular space is on the outside o
the vesicle. !ssume that each pump transports one Na
+
ion in one direction and one K
+

ion in the other direction during each pumping cycle (see 7igure </2=/> or ho2 the
Na
+
$K
+
pump normally unctions in the plasma membrane).
7igure /2=/>
)redict 2hat 2ould happen in each o the ollo2ing conditions-
!. The solutions inside and outside the vesicles contain both Na
+
and K
+
ions but no
!T).
#. The solution outside the vesicles contains both Na
+
and K
+
ions8 the solution
inside contains both Na
+
and K
+
ions and !T).
C. The solution outside contains Na
+
8 the solution inside contains Na
+
and !T).
1&(
12-16 7or each o the ollo2ing sentences+ ill in the blanks 2ith the best 2ord or phrase
selected rom the list belo2. Not all 2ords or phrases 2ill be used8 each 2ord or phrase
should be used only once.
7or an uncharged molecule+ the direction o passive transport across a
membrane is determined solely by its 999999999999999999 gradient. 'n
the other hand+ or a charged molecule+ an additional orce called the
999999999999999999 must also be considered. The net driving orce or
a charged molecule across a membrane thereore has t2o components and
is reerred to as the 999999999999999999 gradient. !ctive transport
allo2s the movement o solutes against this gradient. The carrier proteins
called 999999999999999999 transporters utili"e the movement o one
solute do2n its gradient to provide the energy to drive the uphill transport
o a second gradient. 3hen this transporter moves both ions in the same
direction across the membrane+ it is considered a(n)
9999999999999999998 i the ions move in opposite directions+ the
transporter is considered a(n) 999999999999999999.
antiport coupled membrane potential
!T) hydrolysis electrochemical symport
concentration light=driven uniport
)!n Channels and $he Me#brane P!$en$ial
12-17 6on channels
(a) only open in response to a signal o some kind.
(b) re:uire input o energy in order to unction.
(c) have no limit to the rate at 2hich they can transport ions.
(d) can transport both negative and positive ions through the same channel.
(e) allo2 passage o ions in both directions.
12-18 ! gated ion channel
(a) stays continuously open 2hen stimulated.
(b) opens more re:uently in response to a given stimulus.
(c) opens more 2idely the stronger the stimulus.
(d) remains closed i unstimulated.
1&4
12-19 ;ou have patch=clamped a single voltage=gated ion channel in a membrane and have
obtained the ollo2ing recording (see 7igure </2=/D!).
E7igure </2=/D!F
7or this channel+ match the recordings depicted in 7igure </2=/D# 2ith the appropriate
choices / through G.
/. ! channel that is closed all the time.
2. ! channel that is open all the time.
0. ! channel 2here the ion lo2 has been reversed.
G. ! channel spending more time in the open coniguration.
E7igure </2=/D#F
1&*
12-20 7or each o the ollo2ing sentences+ ill in the blank 2ith the appropriate type o gating
or the ion channel described. ;ou can use the same type o gating mechanism more than
once.
!. The acetylcholine receptor in skeletal muscle cells is a 999999999999999999 ion
channel.
#. 999999999999999999 ion channels are ound in the hair cells o the mammalian
cochlea.
C. 999999999999999999 ion channels in the mimosa plant propagate the lea=
closing response.
&. 999999999999999999 ion channels respond to changes in membrane potential.
(. Hany receptors or neurotransmitters are 999999999999999999 ion channels.
12-21 The high intracellular concentration o K
+
in a resting animal cell is partly due to
(a) the K
+
leak channels in the plasma membrane.
(b) the Na
+
$K
+
pump in the plasma membrane.
(c) voltage=gated K
+
channels in the plasma membrane.
(d) intracellular stores o K
+
in the endoplasmic reticulum.
(e) the membrane potential.
12-22 Itate 2hether you agree or disagree 2ith the ollo2ing statement. (,plain your ans2er.
The membrane potential arises rom movements o charge that leave ion
concentrations practically unaected and result in only a very slight
discrepancy in the number o positive and negative ions on the t2o sides
o the membrane.
)!n Channels and Signaling in Ner+e Cells
12-23 Hatch the numbered lines 2ith the ollo2ing structures-
!. Nerve terminal
#. Cell body
C. !,on
&. &endrite
7igure </2=20
1&,
12-24 7or each o the ollo2ing sentences+ ill in the blanks 2ith the best 2ord or phrase
selected rom the list belo2. Not all 2ords or phrases 2ill be used8 each 2ord or phrase
should be used only once.
The action potential is a 2ave o 999999999999999999 that rapidly
spreads along the neuronal plasma membrane. This 2ave is triggered by a
local change in the membrane potential to a value that is
999999999999999999 negative than the resting membrane potential. The
action potential is propagated by the opening o 999999999999999999=
gated channels. &uring an action potential+ the membrane potential
changes rom 999999999999999999 to 999999999999999999. The action
potential travels rom the neuron4s 999999999999999999 along the
999999999999999999 to the nerve terminals. Neurons chiely receive
signals at their highly branched 999999999999999999.
anions depolari"ation negative
a,on hyperpolari"ation neutral
cell body less positive
cytoskeleton ligand pressure
dendrites more voltage
12-25 'n the diagram o an action potential sho2n belo2+ dra2 a dashed line sho2ing 2hat
2ould have happened to the membrane potential ater the initial depolari"ing stimulus i
there had been no voltage=gated Na
+
channels in the membrane.
7igure </2=2>
1&-
12-26 3hen a neuron receives a signal+ an action potential can be triggered. ! graph o the
traveling membrane depolari"ation assumes a characteristic shape as the depolari"ation
moves do2n the a,on (i.e.+ the peak height and length o depolari"ation is constant).
3hat property o the voltage=gated Na
+
channel ensures that the action potential assumes
this characteristic orm5 (,plain your ans2er.
12-27 7or each o the ollo2ing sentences+ ill in the blanks 2ith the best 2ord or phrase
selected rom the list belo2. Not all 2ords or phrases 2ill be used8 each 2ord or phrase
should be used only once.
Neurons communicate 2ith each other through speciali"ed sites called
999999999999999999. Hany neurotransmitter receptors are ligand=gated
ion channels that open transiently in the 999999999999999999 cell
membrane in response to neurotransmitters released by the
999999999999999999 cell. Jigand=gated ion channels in nerve cell
membranes convert 999999999999999999 signals into
999999999999999999 ones. Neurotransmitter release is stimulated by the
opening o voltage=gated 999999999999999999 in the nerve terminal
membrane.
acetylcholine receptor ?!#! receptor presynaptic
Ca
2+
channels K
+
channels synapses
chemical Na
+
channels
electrical postsynaptic
12-28 6nhibitory neurotransmitters such as ?!#! oten open Cl
$
channels. (,plain ho2 the
opening o a Cl
$
channel 2ill aect the iring o an action potential.
12-29 K
+
leak channels are important or setting the resting membrane potential. 6 you 2ere to
add a drug that inhibits K
+
leak channels in neurons+ 2ould this make it easier or harder
to trigger an action potential in the treated neurons5 3hy5
1&%
H!. /e 0n!.1 S2uid Re+eal Secre$s !" Me#brane E3ci$abili$4
12-30 Itudies on the s:uid giant a,on 2ere instrumental in our current understanding o ho2
action potentials are generated. ;ou decide to do some e,periments on the s:uid giant
a,on yoursel.
!. ;ou remove the cytoplasm in an a,on and replace it 2ith an artiicial cytoplasm
that contains t2ice the normal concentration o K
+
by adding K'!c+ 2here '!c
$

is an anion that is impermeable to the membrane. 6n this 2ay you double the
internal concentration o K
+
2hile maintaining bulk electrical balance o the
cytoplasmic solution. 3ill this make the resting potential o the membrane more
or less negative5
#. ;ou add NaCl to the e,tracellular luid and eectively double the amount o
e,tracellular Na
+
cation. Ho2 does this aect the action potential5
C. ;ou replace hal o the NaCl in the e,tracellular luid 2ith choline chloride.
(Choline is a monovalent cation much larger than Na
+
. Note that the presence o
choline 2ill not impede the lo2 o Na
+
through its channels.) Ho2 2ill this
aect the action potential5
1&&
Ans.ers
12-1 (d)
12-2 The t2o basic properties governing the likelihood o 2hether a molecule 2ill diuse
through a lipid bilayer are the si"e o the molecule and the charge o the molecule. !
smaller molecule 2ill be more likely to diuse through the lipid bilayer than a larger
molecule. ! nonpolar (hydrophobic) molecule 2ill be more likely to diuse through the
lipid bilayer than a polar molecule+ 2hich is more likely to diuse through the lipid
bilayer than a charged molecule.
!. ben"ene (small nonpolar vs. larger uncharged)
#. ethanol (polar vs. charged)
C. glucose (large polar vs. very large highly charged)
&. '2 (nonpolar vs. polar)
(. adenosine (polar vs. highly charged)
12-3 Choice (d) is the correct ans2er. #ecause the lipid bilayer is permeable to carbon dio,ide
and ethanol+ destroying membrane proteins is unlikely to aect their e,it (choices (a) and
(b)). The lipid bilayer is also permeable to 2ater (choice (e)). 'n the other hand glucose
re:uires a membrane transport protein to be imported into the cell. !T)+ 2hich is a highly
charged molecule+ also re:uires a transport protein to cross a membrane and thus could
not be lost rom the cell by simple diusion (choice (c)).
12-4 ! molecule moves do2n its concentration gradient by passive transport+ but re:uires
active transport to move up its concentration gradient. Carrier proteins and ion channels
unction in membrane transport by providing a hydrophilic path2ay through the
membrane or speciic polar solutes or inorganic ions. Carrier proteins are highly
selective in the solutes they transport+ binding the solute at a speciic site and changing its
conormation in order to transport the solute across the membrane. 'n the other hand+
ion channels discriminate among solutes mainly on the basis o si"e and electrical
charge.
200
12-5 !. Holecule # is more likely to utili"e a carrier protein. 7rom the graph+ the yeast
cell4s uptake o molecule # levels o 2hen the concentration o carbon source is
high8 this indicates a saturation point or the uptake o molecule #. Carrier
proteins 2ork by speciically binding their solutes and transerring the solute
molecules across the membrane one at a time+ do2n the concentration gradient o
the solute. Thus+ the rate at 2hich a solute can be transported by a carrier protein
is limited by the number o carrier proteins in the membrane and 2ill reach a
ma,imum 2hen the solute concentration is high enough or solute molecules to
saturate the carrier proteins. Iince the graph indicates that the uptake o molecule
# reaches a saturation point at high concentrations+ molecule # is more likely to
utili"e a carrier protein.
#. Holecule ! is more likely to be ethanol. 1ptake o molecule ! is proportional to
its concentration+ 2hile the uptake o molecule # reaches a saturation point at
high concentration. The linear dependence o molecule ! upon its concentration
suggests that molecule ! can diuse into the cell. Iince ethanol (a small polar
molecule) is more likely to diuse into the cell compared to acetate (a charged
molecule)+ ethanol is likely to be molecule !. Iince acetate is charged+ it is more
likely to re:uire a carrier protein in order to be transported across the membrane.
12-6 !T)=driven transport+ coupled transport+ light=driven transport
12-7 (a)+ (b)+ and (e)
12-8 (d) 6 the pump is mechanistically similar to the Na
+
$K
+
pump+ then the transport o
ions is driven by !T) hydrolysis+ the pump is transiently phosphorylated8
phosphorylation is stimulated by one ion and dephosphorylation is stimulated by
the other ion. Iince all o the protein is in the phosphorylated orm in the absence
o @n
2+
(lane 7)+ @n
2+
is probably re:uired or dephosphorylation. K
+
+ then+
probably binds to the dephosphorylated orm and stimulates the
!T)aseKautophosphorylation. Io+ i @n
2+
is added to the phosphorylated pump+
@n
2+
2ill stimulate dephosphorylation+ trigger a conormational change+ and be
in*ected into the vesicle. K
+
2ill stimulate the kinase activity o the pump+ but
since there is no !T) to be hydroly"ed in the interior o the vesicle+ no
phosphorylation and hence no movement o K
+
2ill occur.
12-9 Na
+
is commonly used to drive coupled transport in animal cells because a steep
concentration gradient o Na
+
(high outside and lo2 inside) is maintained by the Na
+
$K
+

pump. Na
+
readily lo2s back into the cell do2n this gradient because o the negative
membrane potential. The energy provided by the lo2 o Na
+
do2n this steep
electrochemical gradient can be harnessed by coupled transporters.
12-10 (d) 'uabain inhibits the Na
+
$K
+
pump and+ thereore+ leads to an increase in the
intracellular concentration o Na
+
+ 2hich is continually leaking into the cell.
1ptake o glucose into epithelial cells occurs via a Na
+
=glucose symport+ 2hich
uses the Na
+
gradient to drive movement o glucose into the cell.
201
12-11 Choice (e) is the correct ans2er. The Na
+
$K
+
pump keeps Na
+

out directly by pumping it
out and keeps Cl
$

out indirectly by helping to maintain the negative membrane potential.
Cells do not have pumps or moving 2ater molecules across the membrane (choice (a))+
since the lipid bilayer is permeable to 2ater. #acteria do not have Na
+
$K
+
pumps in their
plasma membranes (choice (b)). The Na
+
$K
+
pump cannot directly remove 2ater
molecules rom the cell8 it helps maintain osmotic balance by pumping out the Na
+
that
leaks in+ 2hich 2ould not help i the cell is in a solution o very lo2 ionic strength
(choice (c)). The plant cell 2all is permeable to 2ater and thereore cannot prevent
osmosis (choice (d)).
12-12 (a)
12-13 Choice (b) is the correct ans2er. The ma*or purpose o the Ca
2+
pumps is to keep the
cytosolic concentration o Ca
2+
lo2. 3hen Ca
2+
does move into the cytosol+ it alters the
behavior o many proteins8 hence Ca
2+
is a po2erul signaling molecule. 6t is not involved
in the catalytic activity o (% en"ymes (choice (c)). Iince the levels o Ca
2+
are very lo2
relative to the levels o K
+
and Na
+
+ the Ca
2+
gradient does not have a signiicant eect on
the osmotic balance o the cell (choice (a)) or the membrane potential (choice (d)). They
are not involved in K
+
import (choice (e)).
12-14 &isagree. ! symporter unctions by transporting t2o solutes in the same direction 2hile
an antiporter transports t2o dierent solutes to opposite sides o the membrane.
Thereore+ lipping the symporter around 2ould not change its characteristics into that o
an antiporter+ as it should still transport both solutes in the same direction.
12-15 !. 3ithout any !T) to provide energy or the Na
+
$K
+
pumps+ no ions 2ill be
pumped.
#. The pumps 2ill utili"e the energy rom !T) hydrolysis to transport Na
+
out o the
vesicles and K
+
into the vesicles. (The pumps 2ill stop 2orking 2hen either the
amount o !T) inside the vesicle is depleted or+ 2hen the K
+
outside o the
vesicles is depleted.)
C. The pump 2ill bind a molecule o Na
+
+ causing the !T)ase activity to hydroly"e
!T) and transer the phosphate group onto the pump. ! conormational change
2ill occur+ leading to release o Na
+
rom the vesicle. Ho2ever+ since there is no
K
+
outside o the vesicle+ the pump 2ill get stuck at that step and subse:uent steps
o the cycle 2ill not occur.
202
12-16 7or an uncharged molecule+ the direction o passive transport across a membrane is
determined solely by its concentration gradient. 'n the other hand+ or a charged
molecule+ an additional orce called the membrane potential must also be considered.
The net driving orce or a charged molecule across a membrane thereore has t2o
components and is reerred to as the electrochemical gradient. !ctive transport allo2s
the movement o solutes against this gradient. The carrier proteins called coupled
transporters utili"e the movement o one solute do2n its gradient to provide the energy to
drive the uphill transport o a second gradient. 3hen this transporter moves both ions in
the same direction across the membrane+ it is considered a(n) symport8 i the ions move
in opposite directions+ the transporter is considered a(n) antiport.
12-17 Choice (e) is the correct ans2er. 6ons can pass either 2ay through a channel8 the direction
in 2hich lo2 takes place depends on the concentration gradient and the membrane
potential. Iome ion channels re:uire a speciic stimulus to open them8 others+ such as K
+

leak channels+ do not (choice (a)). 6on channels are passive transporters and thereore
re:uire no energy source in order to unction (choice (b)). 6on channels are very ast
relative to carrier proteins but are limited by the rate at 2hich ions can move through the
channel (choice (c)). !n ion channel allo2s speciic positive or negative ions to pass+ but
not both (choice (d)).
12-18 (b)
12-19 /Lc8 2Ld8 0Lb8 GLa
12-20 !. The acetylcholine receptor in skeletal muscle cells is a liand-ated ion channel.
#. !tress-activated ion channels are ound in the hair cells o the mammalian
cochlea.
C. "oltae-ated ion channels in the mimosa plant propagate the lea=closing
response.
&. "oltae-ated ion channels respond to changes in membrane potential.
(. Hany receptors or neurotransmitters are liand-ated ion channels.
12-21 (b)+ (e) The Na
+
$K
+
pump continually transports K
+
into the cell. The negative membrane
potential also helps to retain K
+
in the cell. The K
+
leak channels allo2 K
+
to
move both into and out o the cell and so do not contribute to the accumulation o
K
+
in the cell.
12-22 !gree. The membrane potential arises rom a thin layer o ions that are close to the
membrane. 'nly a small number o ions (compared to the number o ions present) must
move across the membrane to set up the membrane potential.
12-23 !LG8 #L/8 CL08 &L2
20(
12-24 The action potential is a 2ave o depolari#ation that rapidly spreads along the neuronal
plasma membrane. This 2ave is triggered by a local change in the membrane potential to
a value that is less negative than the resting membrane potential. The action potential is
propagated by the opening o voltae=gated channels. &uring an action potential+ the
membrane potential changes rom neative to positive. The action potential travels rom
the neuron4s cell body along the a$on to the nerve terminals. Neurons chiely receive
signals at their highly branched dendrites.
12-25 Iee igure !/2=2>.
7igure !/2=2>
12-26 The ability o the voltage=gated Na
+
channel to adopt an inactivated conormation allo2s
or the action potential to move along the membrane in a 2ave=like ashion. &uring the
iring o an action potential+ the voltage=gated Na
+
channel 2ill open and then+ ater a
delay+ adopt the inactive orm. 3hen the voltage=gated Na
+
channel adopts an inactive
orm+ the membrane potential begins to return to its resting potential and cannot support
the production o another action potential (i.e.+ urther membrane depolari"ation) until the
channel changes rom an inactive to a closed state.
12-27 Neurons communicate 2ith each other through speciali"ed sites called synapses. Hany
neurotransmitter receptors are ligand=gated ion channels that open transiently in the
postsynaptic cell membrane in response to neurotransmitters released by the
presynaptic cell. Jigand=gated ion channels in nerve cell membranes convert chemical
signals into electrical ones. Neurotransmitter release is stimulated by the opening o
voltage=gated Ca
2%
channels in the nerve terminal membrane.
204
12-28 To trigger an action potential+ a threshold level o membrane depolari"ation must be
achieved to allo2 or the opening o voltage=gated Na
+
channels. !s an action potential
is initiated+ the initial opening o voltage=gated Na
+
channels causes Na
+
to enter the cell.
This makes the membrane potential less negative and opens even more voltage=gated Na
+
channels+ creating a positive eedback loop that triggers the action potential. The opening
o Cl
$
channels by binding o inhibitory neurotransmitters permits an inlu, o Cl
$
+ 2hich
makes the membrane potential more negative. The entry o Cl
$
into the cell counteracts
the eect o the incoming Na
+
on the membrane potential+ making it more diicult to
depolari"e the membrane and thereore+ more diicult to reach the threshold potential
needed to generate an action potential.
12-29 The K
+
leak channel inhibitor 2ould make it easier to trigger an action potential in the
neuron. 6nhibition o the K
+
leak channel changes the permeability o the membrane 2ith
respect to K
+
. The membrane potential is aected by the relative permeability o Na
+
and
K
+
. 6n order or an action potential to ire+ a threshold membrane potential must be
achieved. #y changing the permeability o the membrane to K
+
2ith the addition o the
inhibitor+ the membrane is no2 closer to the threshold needed to ire an action potential
because the cell is no2 partially depolari"ed (i.e.+ addition o the inhibitor leads to a less
negative membrane potential). Thereore+ less Na
+
needs to lo2 into the neuron or the
threshold potential to be achieved+ making it easier to trigger an action potential.
12-30 !. 6ncreasing the concentration o K
+
in the s:uid a,on cytoplasm 2ill make the
membrane potential more negative. &oubling the amount o K
+
increases the
driving orce or K
+
to move out o the cell+ leaving the inside o the cell more
negative and thus decreasing the membrane potential. (%emember+ rom the
Nernst e:uation+ the driving orce or an ion across a membrane is proportional to
the ratio o the concentration o the ion on the outside to the concentration o the
ion on the inside.)
#. &oubling the amount o Na
+
in the e,tracellular luid 2ill increase the height o
the peak o the action potential. !gain+ this is because no2 the driving orce or
Na
+
to enter the cell is greater than it 2as beore. Thus+ 2hen Na
+
channels open+
the lu, o Na
+
ions is no2 greater. (%emember that lu, is the number o ions
entering per second.)
C. The action potential in this case 2ill reach a height that is less than that normally
achieved. (Choline is added in this case to maintain bulk electrical neutrality.
Iince Na
+
channels are not permeable to choline+ they do not contribute to the
electrochemical gradient.) ;ou have no2 reduced the concentration o Na
+
by one
hal and thus decreased the driving orce or Na
+
to enter the cell.
20*