Muscle 381
Figure 12.25 A comparison of the rates at which maximum tension is developed in three muscles. These are ( a ) the
relatively fast-twitch extraocular and ( b ) gastrocnemius muscles, and ( c ) the slow-twitch soleus muscle.
ab c
Time (msec)
Tension
abilities. In some animals, the extreme fast glycolytic fibers are
of the type designated type IIB fibers. In humans, these fast
glycolytic fibers have a different myosin protein and an even
lower oxidative ability than the type IIB fibers, and so are des-
ignated as type IIX fibers. During maximal exercise, the type
IIX fibers have the greatest rate of ATP consumption and the
fastest rate of ATP and phosphocreatine depletion. The three
major fiber types in humans are compared in table 12.3.
People vary tremendously in the proportion of fast- and
slow-twitch fibers in their muscles ( fig. 12.26 ). The percentage
of slow-twitch, type I fibers in the quadriceps femoris muscles
of the legs, for example, can vary from under 20% (in people
who are excellent sprinters) to as high as 95% (in people who
are good marathon runners). These differences are believed to
primarily result from differences in genetics, although physical
training is also an important factor. This is demonstrated by the
complete disappearance of type 1 fibers from human muscle
after long-term spinal cord injury ( fig. 12.26 ).
A muscle such as the gastrocnemius contains both fast and
slow-twitch fibers, although fast-twitch fibers predominate.
A given somatic motor axon, however, innervates muscle fibers
of one type only. The sizes of these motor units differ; the
motor units composed of slow-twitch fibers tend to be smaller
(have fewer fibers) than the motor units of fast-twitch fibers.
As mentioned earlier, motor units are recruited from smaller
to larger when increasing effort is required; thus, the smaller
motor units with slow-twitch fibers would be used most often
in routine activities. Larger motor units with fast-twitch fibers,
which can exert a great deal of force but which metabolize
anaerobically and thus fatigue quickly, would be used rela-
tively infrequently and for only short periods of time.
Muscle Fatigue
Muscle fatigue may be defined as a reversible, exercise-
induced reduction in the ability of a muscle to generate force.
Fatigue during a sustained maximal contraction, when all the
motor units are used and the rate of neural firing is maximal—
as when lifting an extremely heavy weight—appears to be due
to an accumulation of extracellular K^1. Concentrations of K^1
Feature Slow Oxidative/Type I (Red) Fast Oxidative/Type IIA (Red) Fast Glycolytic/Type IIX (White)
Diameter Small Intermediate Large
Z-line thickness Wide Intermediate Narrow
Glycogen content Low Intermediate High
Resistance to fatigue High Intermediate Low
Capillaries Many Many Few
Myoglobin content High High Low
Respiration Aerobic Aerobic Anaerobic
Oxidative capacity High High Low
Glycolytic ability Low High High
Twitch rate Slow Faster Fastest
Myosin ATPase rate Low Higher Highest
Table 12.3 | Characteristics of Muscle Fiber Types