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Essential Cell Biology 5th edition

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Activated Carriers and Biosynthesis

109

TABLE 3–2 SOME ACTIVATED CARRIERS WIDELY USED IN METABOLISM

Activated Carrier

Group Carried in High-Energy Linkage

ATP

phosphate

NADH, NADPH, FADH 2

electrons and hydrogens

Acetyl CoA

acetyl group

Carboxylated biotin

carboxyl group

S-adenosylmethionine

methyl group

Uridine diphosphate glucose

glucose

In acetyl CoA and the other activated carriers in Table 3−2, the transferable

group makes up only a small part of the molecule. The rest consists

of a large organic portion that serves as a convenient “handle,” facilitating

the recognition of the carrier molecule by specific enzymes. As with

acetyl CoA, this handle portion very often contains a nucleotide. This

curious fact may be a relic from an early stage of cell evolution. It is

thought that the main catalysts for early life-forms on Earth were RNA

molecules (or their close relatives) and that proteins were a later evolutionary

addition. It is therefore tempting to speculate that many of the

activated carriers that we find today originated in an earlier RNA world,

where their nucleotide portions would have been useful for binding these

carriers to RNA-based catalysts, or ribozymes (discussed in Chapter 7).

Activated carriers are usually generated in reactions coupled to ATP

hydrolysis, as shown for biotin in Figure 3–38. Therefore, the energy that

enables their groups to be used for biosynthesis ultimately comes from

the catabolic reactions that generate ATP. The same principle applies to

the synthesis of large macromolecules—nucleic acids, proteins, and polysaccharides—as

we discuss next.

acetyl

group

nucleotide

ADENINE

H 3 C

C

O

acetyl group

S

high-energy

bond

H

C

H

H

C

H

O H

N C C

H H

H

C

H

O H CH 3 H O

N C C C C O P O

H OH CH 3 H O –

coenzyme A (CoA)

O

P

O –

O CH 2

RIBOSE

O

– O P O

O –

Figure 3–37 Acetyl coenzyme A (CoA) is

another important activated carrier.

A ball-and-stick model is shown above the

structure of acetyl CoA. The sulfur atom

(orange) forms a thioester bond to acetate.

Because the thioester bond is a high-energy

linkage, it releases a large amount of free

energy when it is hydrolyzed. Thus the

acetyl group carried by CoA can be readily

transferred to other molecules.

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