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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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CHLOROPLASTS AND PHOTOSYNTHESIS

783

LEAF

upper epidermis

chloroplast

grana

thylakoids

air space

nucleus

vacuole

cell wall

cell

wall

lower epidermis

cytosol

chloroplast

mitochondrion

chloroplast

envelope

(A)

(B) (C) (D)

10 mm

5 µm

0.5 µm

not folded into cristae and does not contain electron-transport chains. Instead,

the electron-transport chains, photosynthetic light-capturing systems, and ATP

synthase are all contained in the thylakoid membrane, a separate, distinct membrane

that forms a set of flattened, disc-like sacs, the thylakoids. The thylakoid

membrane is highly folded into numerous local stacks of flattened vesicles called

MBoC6 n14.318,14.34/14.37

grana, interconnected by nonstacked thylakoids. The lumen of each thylakoid is

connected with the lumen of other thylakoids, thereby defining a third internal

compartment called the thylakoid space. This space represents a separate compartment

in each chloroplast that is not connected to either the intermembrane

space or the stroma.

Chloroplasts Capture Energy from Sunlight and Use It to Fix

Carbon

We can group the reactions that occur during photosynthesis in chloroplasts into

two broad categories:

1. The photosynthetic electron-transfer reactions (also called the “light

reactions”) occur in two large protein complexes, called reaction centers,

embedded in the thylakoid membrane. A photon (a quantum of light)

knocks an electron out of the green pigment molecule chlorophyll in the

first reaction center, creating a positively charged chlorophyll ion. This

electron then moves along an electron-transport chain and through a second

reaction center in much the same way that an electron moves along

the respiratory chain in mitochondria. During this electron-transport

process, H + is pumped across the thylakoid membrane, and the resulting

Figure 14–37 Chloroplasts in the cell.

(A) Schematic cross section through the

leaf of a green plant. (B) Light microscopy

of a plant leaf cell—here, a mesophyll cell

from Zinnia elegans—shows chloroplasts

as bright green bodies, measuring several

micrometers across, in the transparent

cell interior. (C) The electron micrograph

of a thin, stained section through a wheat

leaf cell shows a thin rim of cytoplasm—

containing chloroplasts, the nucleus, and

mitochondria—surrounding a large, waterfilled

vacuole. (D) At higher magnification,

electron microscopy reveals the chloroplast

envelope membrane and the thylakoid

membrane within the chloroplast that

is highly folded into grana stacks

(Movie 14.9). (B, courtesy of John Innes

Foundation; C and D, courtesy of

K. Plaskitt.)

thylakoid membrane

inner membrane

outer membrane

intermembrane space

matrix stroma

thylakoid space

crista

grana

crista membrane thylakoids

2 µm

MITOCHONDRION

CHLOROPLAST

Figure 14–38 A mitochondrion and

chloroplast compared. Chloroplasts

are generally larger than mitochondria. In

addition to an outer and inner envelope

membrane, they contain the thylakoid

membrane with its internal thylakoid space.

The chloroplast thylakoid membrane,

which is the site of solar energy conversion

in plants and algae, corresponds to the

mitochondrial cristae, which are the sites

of energy conversion by cellular respiration.

Unlike the crista membrane, which is

continuous with the inner mitochondrial

membrane at cristae junctions, the thylakoid

membrane is not connected to the inner

chloroplast membrane at any point.

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