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Chapter 11 - Voltages and Free Energy

True/False and Multiple Choice (7)


The oxidative half-reaction involves the release of electrons from a molecule



Which of the following is a common consequence of a protein coordinating a

a) the free energy of the reduced and oxidized form of the metal are altered
b) the metal is protected from some chemical reactions
c) the protein undergoes a conformational change
d) it enables the transfer of electrons over long distances
e) all of the above


Standard reduction potentials in biochemistry are pH independent



Which of the following conformational changes is not thought to occur as a K+

channel closes due to a voltage change:
a) Helix S6 becomes straighter
b) Helix S4 is moves downward
c) Charged residues on S4 form new interactions
d) The selectivity filter changes size
e) The C-terminus of S4 forms a 3-10 alpha helix


Most ion channels have little selectivity because the sizes of ions is small relative
to the size of the channel pore.


Which feature allows mitochondria and chloroplasts, unique from other

organelles, to maintain large proton gradients:
a) they contains membrane spanning proteins
b) they has a second internal membrane
c) there are only one of each per cell
d) their membranes consists of a lipid bilayer


Which of the following statements about passive spread are true:

I - the voltage spike is not regenerated
II - the potassium conductance of the membrane changes locally
III - the sodium conductance of the membrane remains constant
IV - it only occurs in non-myelinated axons
a) none of the above
b) all of the above
c) I and III
d) I, II and III
e) II and IV

Fill in the Blank (5)


The oxidized and reduced forms of a molecule are known as a ____________

redox couple


_______ is used to reduce metals in photosynthesis.



Sodium and Potassium pumps hydrolyze ________ to move ions in and out of
the cell


By increasing the capacitance of the surface of the axon, ______________

greatly facilitates the transmission of action potentials


A _________________ can build up charge on its surface when sandwiched

between two conductors.

Long Answer/Essay (12)


How does phosphorylation contribute to increasing the diversity of redox active

molecules in the cell?
Organic compounds that undergo redox reactions can be phosphorylated on
parts of the molecule that are not important for the redox reaction. For example,
NAD+/NADH can be converted to NADP+/NADPH. The phosphorylated and
unphosphorylated forms of the compound bind specifically to different subsets of
cellular proteins, which increase the diversity of potentially active redox molecules
in the cell.


A new microrganism is isolated from a lake and is placed into a solution of KCl.
The voltage difference across its membrane is measured at 120 mV. How much
energy is required to move a proton from the negative side of the membrane to the
positive side?
U = q x E
= 96500 C/mol x 0.12 J/C
= 11580 J/mol
= 11.58 kJ/mol


An electrochemical cell couples manganese and copper. a) What is the

combined redox reaction? b) What is the G for the reaction?

a) Mn -> Mn2+ + 2e-, E = 1.18 V (positive, the standard reaction is reversed)

2e- + Cu2+ -> Cu, E = 0.340 V
Cu2+ + Mn -> Mn2+ + Cu, E= 1.52 V
b) G = -FE
= -2 x 96500C/mol x 1.52 J/C
= -293360 J
= -293.4 kJ/mol


A researcher assembles an electrochemical cell with silver and an unknown

metal. She measures the free energy of the combined reaction under standard
conditions to be -150 kJ/mol. Assuming standard conditions, what is the potential of
the oxidative half reaction if one electron is transferred by the unknown metal to
E = - G/F = 150000J / 96500C/mol * 1
= 1.55 V
Eunknown = E - EAg = 1.55 V - 0.8 V = 0.75 V


A new ATP-producing protein is discovered that takes couples ATP production to

the oxidation of NADPH by oxidative phosphorylation. If this protein only produces 1
molecule of ATP per reaction that consumes one NADPH: a) how much energy is
wasted? b) how many more ATP molecules could be created by a perfectly efficient
electron transport chain from one NADPH?
(assume the G for ATP synthesis is 30kJ/mol)
a) The reaction:
NADPH -> NADP+ + H+ + 2e-, E = 0.32
1/2 O2 + 2H+ + 2e- -> H2O, E = 0.815
NADPH + H+ + 1/2 O2 -> H2O + NADP+, E = 1.135
G = -FE = - 2 x 96500 C/mol x 1.135 J/C
= 219 kJ/mol
Gwasted = G - GATP = 219 kJ - 30 kJ = 189 kJ
b) 189/30 = 6.3 more ATPs could be produced.


Consider the Potassium channel: a) Given that the four K+ binding sites are
roughly isoenergetic, why are usually only two occupied at any given time? b) How
does this contribute to the knock on model for conductance? c) What establishes
the directionality of K+ ion flow?

a) The binding sites are close enough such that electrostatic repulsion of K+ ions
in adjacent sites is highly unfavourable. In contrast, occupancy of alternating
binding sites (1 and 3, or 2 and 4) is favourable.
b) Alternating binding sites means that when something enters site if sites 2 and
4 are occupied, binding at site 1, will move an ion from 2 to 3, and from 4 into
the vestibule. Alternatively, if sites 1 and 3 are occupied, binding at site 4 will
move an ion from 3 to 2, and from 1 into the extracellular space
c) the directionality is established by the electrochemical gradient. There is no
directionality preference for the channel itself.

A scientist measures the potential across a lipid bilayer. At room temperature,

the pH outside is 7.4 and inside the pH is 7.1. What is the potential for protons
across the bilayer?
EH = RT/F x ln ([H+]out/[H+]in) = 59mV x 7.4/7.1 = 61.5 mV


A squid is immersed seawater where its neurons are at a resting potential of -76
mV a) What are the potentials for Na and K in seawater? b) Why is the resting
potential not equal to the Na, K or sum of potentials?







a) EK = 0.0257V x ln ([K+]out/[K+]in) = 25.7mV x -3.4 = -87.4 mV
ENa = 0.0257V x ln ([Na+]out/[Na+]in) = 25.7mV x 1.9 = 48.8 mV
b) The resting potential is influenced by the relative conductance of the two ions
across the membrane. Since the membrane is more permeable to K+ than
Na+, the actual potential is weighted more heavily to the K+ potential (see also
equation 11.76)


A scientist grows a tiny synthetic nerve cell with a surface area of 3.1 x 10 -7 cm2.
In its culture medium, the cell has a resting potential of 75 mV. If 6.2 pC of charge
move across the membrane during an action potential that depolarizes the cell to 0
mV. What is the capacitance of the cell per unit area of the cell?
q = CxV
C = V/q = 75mV/6.2 pC = 0.075V/6.2 x 10-12 C
= 4.65 10-13 F
C = CA x A
CA = C/A = 4.65 10-13 F/ 3.1 x 10-7 cm2
= 1.5 x 10-6 F/cm = 1.5 uF / cm2


A cell has the intracellular and extracellular concentrations of K+ and Na+ listed
below. If the resting conductance of its K+ Channels is 30 times greater than its
Na+ Channels, what is the resting potential of the cell (assuming no other ions
contribute significantly)?








E = (gNa/(gNa+ gK))xENa + (gK/(gNa+ gK))xEK
= (1/(1+30))x 0.0257V x ln (360/12) + (30/(1+30))x 0.0257V x ln (200/15)
= 2.8mV - 64.4 mV
= -61.6 mV
A region of an axon has a time constant of 120 msec. The capacitance of the
membrane is 1 uF/cm2 and there are 25000 potassium channels/cm2. How many ions
does a typical potassium channel conduct per second against a potential of 100mV?

First solve the resistance:

t = RK x C
RK = C/t = 1uF/cm2 / 0.12s = 120000 ohm/cm2
Next, find the current:
IM = EM/RK = 0.1V/120000 ohm/cm2
= 8.3 x 10-7 C/s/cm2
Finally, solve for flow:
IM = Flow * F * #channels/NA
Flow = IM*NA/F*channels
= (8.3 x 10-7 C/s/cm2 * 6.023x1023 ) / ( 96500 * 25000)
= 2.07 x 108 C per second
Immediately prior and subsequent to an action potential, there is little
conductance through sodium channels; however, conductance is blocked through
different underlying mechanisms. What forces drive the conformational features that
distinguish the closed and inactivated states of the Na channel?
Prior to the action potential, the voltage sensor is in the closed conformation,
which is established by interactions of the positively charged residues on the
voltage sensor and the negative electric field on the inside of the membrane.
This causes the helices that comprise the channel to be closed.
After the action potential, the electric field is not as negative, so the voltage
sensor is in the open conformation, but flexible cytosolic inactivation domains
bind to the base of the channel and prevent ion conductance.