100 Chapter 4
a particular atom or molecule can play both roles; it may func-
tion as an oxidizing agent in one reaction and as a reducing agent
in another reaction. When atoms or molecules play both roles,
they gain electrons in one reaction and pass them on in another
reaction to produce a series of coupled oxidation-reduction
reactions—like a bucket brigade, with electrons in the buckets.
Notice that the term oxidation does not imply that oxygen
participates in the reaction. This term is derived from the fact
that oxygen has a great tendency to accept electrons; that is,
to act as a strong oxidizing agent. This property of oxygen is
exploited by cells; oxygen acts as the final electron acceptor in
a chain of oxidation-reduction reactions that provides energy
for ATP production (chapter 5, section 5.2).
Oxidation-reduction reactions in cells often involve the
transfer of hydrogen atoms rather than free electrons. Because a
hydrogen atom contains 1 electron (and 1 proton in the nucleus),
a molecule that loses hydrogen becomes oxidized, and one that
gains hydrogen becomes reduced. In many oxidation-reduction
reactions, pairs of electrons—either as free electrons or as a pair
of hydrogen atoms—are transferred from the reducing agent to
the oxidizing agent.
Two molecules that serve important roles in the transfer
of hydrogens are nicotinamide adenine dinucleotide (NAD),
which is derived from the vitamin niacin (vitamin B 3 ), and flavin
adenine dinucleotide (FAD), which is derived from the vitamin
riboflavin (vitamin B 2 ). These molecules ( fig. 4.17 ) are coen-
zymes that function as hydrogen carriers because they accept
hydrogens (becoming reduced) in one enzyme reaction and
donate hydrogens (becoming oxidized) in a different enzyme
reaction ( fig. 4.18 ). The oxidized forms of these molecules are
written simply as NAD (or NAD^1 ) and FAD.
Figure 4.16 A model of ATP as the universal energy carrier of the cell. Exergonic reactions are shown as blue gears
with arrows going down (these reactions produce a decrease in free energy); endergonic reactions are shown as green gears with
arrows going up (these reactions produce an increase in free energy).
ATP produced ATP used for cell work
Food
AT P
ADP + Pi AT P
ADP + Pi
AT P
ADP + Pi
AT P
ADP + Pi
AT P
ADP + Pi
AT P
ADP + Pi
CO 2 + H 2 O
Figure 4.15 The formation and structure of
adenosine triphosphate (ATP). ATP is the universal energy
carrier of the cell. High-energy bonds are indicated by a squiggle
(~). When the bond to the last phosphate is broken, energy is
released and ATP is converted back into ADP and P i.
Adenosine diphosphate (ADP)
Inorganic phosphate (Pi)
NH 2
CH 2
C N
C
C
CH
HC
N
N
N
H H
H
OH
Adenosine triphosphate (ATP)
+
H
OOP~ O
OH
_
O
_
O
P~O
O
_
O
OP
O
_
O
donates electrons to another, which therefore becomes reduced.
The atom or molecule that donates electrons to another
is a reducing agent, and the one that accepts electrons from
another is an oxidizing agent. It is important to understand that