110 Chapter 5
Red blood cells lack mitochondria and so can metabolize
only anaerobically; they cannot use the oxygen they carry. This
spares the oxygen within red blood cells for delivery to the
other cells of the body. Except for red blood cells, anaerobic
metabolism occurs for only a limited period of time in tissues
that have energy requirements in excess of their aerobic ability.
Anaerobic metabolism occurs in the skeletal muscles and heart
when the ratio of oxygen supply to oxygen need (related to the
concentration of NADH) falls below a critical level. Anaerobic
metabolism is, in a sense, an emergency procedure that provides
some ATP until the emergency (oxygen deficiency) has passed.
It should be noted, though, that there is no real “emergency”
in the case of skeletal muscles, where lactic acid fermentation
is a normal, daily occurrence that does not harm muscle tis-
sue or the individual. Excessive lactic acid production by mus-
cles, however, is associated with pain and muscle fatigue. (The
metabolism of skeletal muscles is discussed in chapter 12, sec-
tion 12.4.) In contrast to skeletal muscles, the heart normally
respires only aerobically. If anaerobic conditions do occur in
the heart, a potentially dangerous situation may be present.
Figure 5.3 Glycolysis. In glycolysis, 1 glucose is
converted into 2 pyruvic acids in nine separate steps. In addition
to 2 pyruvic acids, the products of glycolysis include 2 NADH
and 4 ATP. Because 2 ATP were used at the beginning, however,
the net gain is 2 ATP per glucose. Dashed arrows indicate
reverse reactions that may occur under other conditions.
Glucose (C 6 H 12 O 6 )
Pyruvic acid (C 3 H 4 O 3 )
ADP
AT P
Glucose 6-phosphate
Fructose 6-phosphate
ADP
AT P
Fructose 1,6-biphosphate
3–Phosphoglyceraldehyde
5
1,3–Biphosphoglyceric acid
Pyruvic acid (C 3 H 4 O 3 )
6
3–Phosphoglyceric acid
7
2–Phosphoglyceric acid
8
Phosphoenolpyruvic acid
9
ADP
AT P
ADP
AT P
ADP
AT P
ADP
AT P
NADH
NAD
2H
NADH
NAD
2H
Pi
3–Phosphoglyceraldehyde
5
1,3–Biphosphoglyceric acid
6
3–Phosphoglyceric acid
7
2–Phosphoglyceric acid
8
Phosphoenolpyruvic acid
9
Pi
Dihydroxy-
acetone
phosphate
1
2
3
4
Figure 5.4 The formation of lactic acid. The addition
of 2 hydrogen atoms (colored boxes) from reduced NAD to
pyruvic acid produces lactic acid and oxidized NAD. This
reaction is catalyzed by lactic acid dehydrogenase (LDH) and is
reversible under the proper conditions.
H C C
HO
H
C
O
OH
O
OH
NADH + H+ NAD
H C C
H
H
C
OH
H
Pyruvic acid Lactic acid
LDH
CLINICAL APPLICATION
Myocardia ischemia refers to inadequate blood flow to the
heart muscle, a condition of ischemic heart disease. It may
also be called coronary heart disease, because of inad-
equate blood flow in the coronary arteries. This is usually
the result of the development of plaques in the artery wall, a
process termed atherosclerosis. Thus, the condition is also
called atherosclerotic heart disease. When the myocar-
dial cells do not obtain sufficient oxygen from the blood for
aerobic respiration, they obtain energy from the less-efficient
lactic acid pathway. Prolonged ischemia can cause death of
these cells, producing myocardial infarction (heart attack).
Lactic acid within the myocardium produces a chest pain
called angina pectoris. Angina is usually treated with vasodi-
lator drugs such as nitroglycerin, which improves blood flow
to the heart and reduces the work of the heart by dilating
peripheral blood vessels.