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Molecular Biology of the Cell by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, Martin Raff, Keith Roberts, Peter Walter by by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morg

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810 Chapter 14: Energy Conversion: Mitochondria and Chloroplasts

the electron carriers in the respiratory chain? What is their

order?

(A)

direction of rotation

14–8 Normally, the flow of electrons to O 2 is tightly

linked to the production of ATP via the electrochemical

gradient. If ATP synthase is inhibited, for example, electrons

do not flow down the electron-transport chain and

respiration ceases. Since the 1940s, several substances—

such as 2,4-dinitrophenol—have been known to uncouple

electron flow from ATP synthesis. Dinitrophenol was

once prescribed as a diet drug to aid in weight loss. How

would an uncoupler of oxidative phosphorylation promote

weight loss? Why do you suppose dinitrophenol is

no longer prescribed?

inner membrane

matrix

(B)

5

β

γ

α

β

actin filament

14–9 In actively respiring liver mitochondria, the pH in

the matrix is about half a pH unit higher than it is in the

cytosol. Assuming that the cytosol is at pH 7 and the matrix

is a sphere with a diameter of 1 μm [V = (4/3)πr 3 ], calculate

the total number of protons in the matrix of a respiring

liver mitochondrion. If the matrix began at pH 7 (equal to

that in the cytosol), how many protons would have to be

pumped out to establish a matrix pH of 7.5 (a difference of

0.5 pH units)?

14–10 ATP synthase is the world’s smallest rotary motor.

Passage of H + ions through the membrane-embedded

portion of ATP synthase (the F o component) causes rotation

of the single, central, axle-like γ subunit inside the

head group. The tripartite head is composed of the three

αβ dimers, the β subunit of which is responsible for synthesis

of ATP. The rotation of the γ subunit induces conformational

changes in the αβ dimers that allow ADP and

Pi to be converted into ATP. A variety of indirect evidence

had suggested rotary catalysis by ATP synthase, but seeing

is believing.

To demonstrate rotary motion, a modified form of

the α 3 β 3 γ complex was used. The β subunits were modified

so they could be firmly anchored to a solid support and the

γ subunit was modified (on the end that normally inserts

into the F o component in the inner membrane) so that a

fluorescently tagged, readily visible filament of actin could

be attached (Figure Q14–2A). This arrangement allows

rotations of the γ subunit to be visualized as revolutions

of the long actin filament. In these experiments, ATP synthase

was studied in the reverse of its normal mechanism

by allowing it to hydrolyze ATP. At low ATP concentrations,

the actin filament was observed to revolve in steps of 120°

and then pause for variable lengths of time, as shown in

Figure Q14–2B.

A. Why does the actin filament revolve in steps with

pauses in between? What does this rotation correspond to

in terms of the structure of the α 3 β 3 γ complex?

B. In its normal mode of operation inside the cell,

how many ATP molecules do you suppose would be synthesized

for each complete 360° rotation of the γ subunit?

Explain your answer.

14–11 How much energy is available in visible light? How

much energy does sunlight deliver to Earth? How efficient

revolutions

4

3

2

1

0

0 20 40 60 80 100

time (seconds)

Figure Q14–2 Experimental set-up for observing rotation of the

γ subunit of ATP synthase (Problem 14–10). (A) The immobilized

α 3 β 3 γ complex. The β subunits are anchored to a solid support and

a fluorescent actin filament is attached to the γ subunit. (B) Stepwise

revolution of the actin filament. The indicated trace is a typical example

Figure p14.04/14.07

from one experiment. The inset shows the positions in the revolution

at which the actin filament pauses. (B, from R. Yasuda et al., Cell

93:1117–1124, 1998. With permission from Problem Elsevier.) p14-31/14-47

are plants at converting light energy into chemical energy?

The answers to these questions provide an important

backdrop to the subject of photosynthesis.

Each quantum or photon of light has energy hv,

where h is Planck’s constant (6.6 × 10 –37 kJ sec/photon)

and v is the frequency in sec –1 . The frequency of light is

equal to c/λ, where c is the speed of light (3.0 × 10 17 nm/

sec) and λ is the wavelength in nm. Thus, the energy (E) of

a photon is

E = hv = hc/λ

A. Calculate the energy of a mole of photons (6 × 10 23

photons/mole) at 400 nm (violet light), at 680 nm (red

light), and at 800 nm (near-infrared light).

B. Bright sunlight strikes Earth at the rate of about 1.3

kJ/sec per square meter. Assuming for the sake of calculation

that sunlight consists of monochromatic light of wavelength

680 nm, how many seconds would it take for a mole

of photons to strike a square meter?

C. Assuming that it takes eight photons to fix one

molecule of CO 2 as carbohydrate under optimal conditions

(8–10 photons is the currently accepted value), calculate

how long it would take a tomato plant with a leaf

area of 1 square meter to make a mole of glucose from CO 2 .

Assume that photons strike the leaf at the rate calculated

above and, furthermore, that all the photons are absorbed

and used to fix CO 2 .

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