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Essential Cell Biology 5th edition

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440 CHAPTER 13 How Cells Obtain Energy from Food

Figure 13–12 The citric acid cycle

catalyzes the complete oxidation of

acetyl groups supplied by acetyl CoA.

The cycle begins with the reaction of

acetyl CoA (derived from pyruvate as

shown in Figure 13–10) with oxaloacetate

to produce citric acid (citrate). The number

of carbon atoms in each intermediate is

shaded in yellow. (See also Panel 13–2,

pp. 442–443.) The steps of the citric acid

cycle are reviewed in Movie 13.2.

NADH

O

H 3 C C S–CoA

acetyl CoA

2C

HS–CoA

oxaloacetate

6C

citrate

4C

STEP 1

STEP 2

6C

STEP 8

NADH

4C

STEP 7

4C

STEP 6

CITRIC ACID

CYCLE

4C

STEP 5

STEP 3

C O 2

5C

STEP 4

4C

NADH

C O 2

FADH 2

GTP

NET RESULT: ONE TURN OF THE CYCLE PRODUCES THREE NADH, ONE GTP, AND

ONE FADH 2 , AND RELEASES TWO MOLECULES OF CO 2

acid cycle are supplied by atmospheric O 2 . In fact, these oxygen atoms

come from water (H 2 O). As illustrated at the top of Panel 13–2, three

molecules of H 2 O are split as they enter each turn of the cycle, and their

oxygen atoms are ultimately used to make CO 2. As we see shortly, the

O 2 that we breathe is actually reduced to H 2 O by the electron-transport

chain; it does not form the CO 2 that we exhale.

Thus far, we have discussed only one of the three types of activated carriers

that are produced by the citric acid cycle—NADH. In addition to three

molecules of NADH, each turn of the cycle also produces one molecule of

FADH 2 (reduced flavin adenine ECB5 e13.12-13.12

dinucleotide) from FAD and one molecule

of the ribonucleoside triphosphate GTP (guanosine triphosphate)

from GDP (see Figure 13–12). The structures of these two activated carriers

are illustrated in Figure 13–13. GTP is a close relative of ATP, and

(A)

– O

O

P

O –

O

O

P

O –

O

O

P

O –

O

O

guanine

N

C

C NH

HC

C C

N N NH

CH 2

2 O

ribose

OH OH

GDP

(B)

H 3 C

H 3 C

C

C

H

C

C

H

2 H + 2 e –

FAD FADH 2

N

O

C

H

N

C C NH

C

C C C

C

N N O

CH 2

N

H

GTP

Figure 13–13 Each turn of the citric acid cycle produces one molecule of GTP

and one molecule of FADH 2 . (A) GTP and GDP are close relatives of ATP and ADP,

respectively, the only difference being the substitution of the base guanine for adenine.

(B) Despite its very different structure, FADH 2 , like NADH and NADPH (see Figure 3–34),

is a carrier of hydrogen atoms and high-energy electrons. It is shown here in its oxidized

form (FAD), with the part of the structure involved in accepting and donating hydrogens

highlighted in yellow. FAD can accept two hydrogen atoms, along with their electrons,

to form the reduced FADH 2 . The atoms involved are shown in their reduced form in the

excerpt to the right.

H

H

H

C

C

C

H 2 C

OH

OH

OH

O P

FAD

P

O

CH 2

RIBOSE

ADENINE

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