320 CHAPTER 10 • MUSCULAR TISSUE
back to glucose. In addition to providing new glucose molecules,
this conversion reduces acidity of the blood. Anaerobic
cellular respiration can provide enough energy for about 30 to
40 seconds of maximal muscle activity. Together, conversion of
creatine phosphate and glycolysis can provide enough ATP to
run a 400-meter race.
Aerobic Cellular Respiration
Muscular activity that lasts longer than half a minute depends
increasingly on aerobic cellular respiration, a series of
oxygen-requiring reactions that produce ATP in mitochondria. If
sufficient oxygen is present, pyruvic acid enters the mitochon-
dria, where it is completely oxidized in reactions that generate
ATP, carbon dioxide, water, and heat (Figure 10.12c). Although
aerobic cellular respiration is slower than glycolysis, it yields
much more ATP. Each molecule of glucose yields about 36 mol-
ecules of ATP; a typical fatty acid molecule yields more than
100 molecules of ATP via aerobic cellular respiration.
Muscle tissue has two sources of oxygen: (1) oxygen that
diffuses into muscle fibers from the blood and (2) oxygen
released by myoglobin within muscle fibers. Both myoglobin
(found only in muscle cells) and hemoglobin (found only in red
blood cells) are oxygen-binding proteins. They bind oxygen
when it is plentiful and release oxygen when it is scarce.
Aerobic cellular respiration supplies enough ATP for pro-
longed activity provided sufficient oxygen and nutrients are avail-
able. These nutrients include the pyruvic acid obtained from the
glycolysis of glucose, fatty acids from the breakdown of triglyc-
erides in adipose cells, and amino acids from the breakdown of
proteins. In activities that last more than 10 minutes, the aerobic
system provides more than 90% of the needed ATP. At the end
of an endurance event such as a marathon race, nearly 100% of
the ATP is being produced by aerobic cellular respiration.
Muscle Fatigue
The inability of a muscle to maintain force of contraction after
prolonged activity is called muscle fatigue. Fatigue results
mainly from changes within muscle fibers. Even before actual
muscle fatigue occurs, a person may have feelings of tiredness
and the desire to cease activity; this response, called central
fatigue, is caused by changes in the central nervous system
(brain and spinal cord). Although its exact mechanism is
unknown, it may be a protective mechanism to stop a person
from exercising before muscles become damaged. As you will
see, certain types of skeletal muscle fibers fatigue more quickly
than others.
Although the precise mechanisms that cause muscle fatigue
are still not clear, several factors are thought to contribute. One
is inadequate release of calcium ions from the SR, resulting in a
decline of Ca2 concentration in the sarcoplasm. Depletion of
creatine phosphate also is associated with fatigue, but surpris-
ingly, the ATP levels in fatigued muscle often are not much
lower than those in resting muscle. Other factors that contribute
to muscle fatigue include insufficient oxygen, depletion of
glycogen and other nutrients, buildup of lactic acid and ADP,
and failure of action potentials in the motor neuron to release
enough acetylcholine.
Oxygen Consumption After Exercise
During prolonged periods of muscle contraction, increases in
breathing rate and blood flow enhance oxygen delivery to
muscle tissue. After muscle contraction has stopped, heavy
breathing continues for a while, and oxygen consumption
remains above the resting level. Depending on the intensity of
the exercise, the recovery period may be just a few minutes, or it
may last as long as several hours. The term oxygen debt refers
to the added oxygen, over and above the resting oxygen con-
sumption, that is taken into the body after exercise. This extra
oxygen is used to “pay back” or restore metabolic conditions to
the resting level in three ways: (1) to convert lactic acid back
into glycogen stores in the liver, (2) to resynthesize creatine
phosphate and ATP in muscle fibers, and (3) to replace the
oxygen removed from myoglobin.
The metabolic changes that occur during exercise can
account for only some of the extra oxygen used after exercise.
Only a small amount of glycogen resynthesis occurs from lactic
acid. Instead, most glycogen is made much later from dietary
carbohydrates. Much of the lactic acid that remains after
exercise is converted back to pyruvic acid and used for ATP
production via aerobic cellular respiration in the heart, liver,
kidneys, and skeletal muscle. Oxygen use after exercise also is
boosted by ongoing changes. First, the elevated body tem-
perature after strenuous exercise increases the rate of chemical
reactions throughout the body. Faster reactions use ATP more
rapidly, and more oxygen is needed to produce the ATP. Second,
the heart and the muscles used in breathing are still working
harder than they were at rest, and thus they consume more ATP.
Third, tissue repair processes are occurring at an increased pace.
For these reasons, recovery oxygen uptake is a better term than
oxygen debt for the elevated use of oxygen after exercise.
CHECKPOINT
11. Which ATP-producing reactions are aerobic and which are
anaerobic?
12. Which sources provide ATP during a 1000-meter run?
13. What factors contribute to muscle fatigue?
14. Why is the term recovery oxygen uptake more accurate
than oxygen debt?
CONTROL OF MUSCLE TENSION
OBJECTIVES
• Describe the structure and function of a motor unit, and
define motor unit recruitment.
• Explain the phases of a twitch contraction.
• Describe how frequency of stimulation affects muscle ten-
sion, and how muscle tone is produced.
• Distinguish between isotonic and isometric contractions.

A single nerve impulse in a somatic motor neuron elicits a single
muscle action potential in all the skeletal muscle fibers with
which it forms synapses. Action potentials always have the same
size in a given neuron or muscle fiber. In contrast, the force of
muscle fiber contraction does vary; a muscle fiber is capable of
producing a much greater force than the one that results from a
single action potential. The total force or tension that a single
muscle fiber can produce depends mainly on the rate at which
nerve impulses arrive at the neuromuscular junction. The num-
ber of impulses per second is the frequency of stimulation.
Maximum tension is also affected by the amount of stretch
before contraction (see Figure 10.9) and by nutrient and oxygen
availability. The total tension a whole muscle can produce
depends on the number of muscle fibers that are contracting
in unison.
Motor Units
Even though each skeletal muscle fiber has only a single neuro-
muscular junction, the axon of a somatic motor neuron branches
out and forms neuromuscular junctions with many different
muscle fibers. A motor unit consists of a somatic motor neuron
plus all the skeletal muscle fibers it stimulates (Figure 10.13). A
single somatic motor neuron makes contact with an average of
150 skeletal muscle fibers, and all of the muscle fibers in one
motor unit contract in unison. Typically, the muscle fibers of
a motor unit are dispersed throughout a muscle rather than
clustered together.
Whole muscles that control precise movements consist of
many small motor units. For instance, muscles of the larynx
(voice box) that control voice production have as few as two or
three muscle fibers per motor unit, and muscles controlling eye
movements may have 10 to 20 muscle fibers per motor unit. In
contrast, skeletal muscles responsible for large-scale and power-
ful movements, such as the biceps brachii muscle in the arm and
Figure 10.13 Motor units. Two somatic motor neurons (one
purple and one green) are shown, each supplying the muscle fibers
of its motor unit.
A motor unit consists of a somatic motor neuron
plus all the muscle fibers it stimulates.
Neuromuscular
junction
Motor
Spinal cord neurons
Muscle fibers (cells)
? What is the effect of the size of a motor unit on its strength
of contraction? (Assume that each muscle fiber can gener-
ate about the same amount of tension.)
CONTROL OF MUSCLE TENSION 321
the gastrocnemius muscle in the calf of the leg, have as many as
2000 to 3000 muscle fibers in some motor units. Because all
the muscle fibers of a motor unit contract and relax together, the
total strength of a contraction depends, in part, on the size of the
motor units and the number that are activated at a given time.
Twitch Contraction
A twitch contraction is the brief contraction of all the muscle
fibers in a motor unit in response to a single action potential in
its motor neuron. In the laboratory, a twitch can be produced by
direct electrical stimulation of a motor neuron or its muscle
fibers. The record of a muscle contraction, called a myogram, is
shown in Figure 10.14. Twitches of skeletal muscle fibers last
anywhere from 20 to 200 msec. This is very long compared to
the brief 1–2 msec* that a muscle action potential lasts.
Note that a brief delay occurs between application of the
stimulus (time zero on the graph) and the beginning of contrac-
tion. The delay, which lasts about two milliseconds, is termed
the latent period. During the latent period, the muscle action
potential sweeps over the sarcolemma and calcium ions are
released from the sarcoplasmic reticulum. The second phase, the
contraction period, lasts 10–100 msec. During this time, Ca2
binds to troponin, myosin-binding sites on actin are exposed,
and crossbridges form. Peak tension develops in the muscle
fiber. During the third phase, the relaxation period, also lasting
10–100 msec, Ca2 is actively transported back into the
sarcoplasmic reticulum, myosin-binding sites are covered by
tropomyosin, myosin heads detach from actin, and tension in the
muscle fiber decreases. The actual duration of these periods
depends on the type of skeletal muscle fiber. Some fibers, such
as the fast-twitch fibers that move the eyes (described shortly),
have contraction periods as brief as 10 msec and equally brief
*One millisecond (msec) is 10-3 seconds (0.001sec).
Figure 10.14 Myogram of a twitch contraction. The arrow
indicates the time at which the stimulus occurred.
A myogram is a record of a muscle contraction.
Contraction
period
Force of contraction
Relaxation
period
Latent
period
0 10 20 30 40 50
Time in milliseconds (msec)
? What events occur during the latent period?
322 CHAPTER 10 • MUSCULAR TISSUE

relaxation periods. Others, such as the slow-twitch fibers that
move the legs, have contraction and relaxation periods of about
100 msec each.
If two stimuli are applied, one immediately after the other,
the muscle will respond to the first stimulus but not to the
second. When a muscle fiber receives enough stimulation to con-
tract, it temporarily loses its excitability and cannot respond for
a time. The period of lost excitability, called the refractory
period, is a characteristic of all muscle and nerve cells. The
duration of the refractory period varies with the muscle
involved. Skeletal muscle has a short refractory period of about
five milliseconds; cardiac muscle has a longer refractory period
of about 300 milliseconds.
Frequency of Stimulation
When a second stimulus occurs after the refractory period of the
first stimulus is over, but before the skeletal muscle fiber has
relaxed, the second contraction will actually be stronger than the
first (Figure 10.15b). This phenomenon, in which stimuli arriv-
ing at different times cause larger contractions, is called wave
summation. When a skeletal muscle fiber is stimulated at a
rate of 20 to 30 times per second, it can only partially relax
between stimuli. The result is a sustained but wavering contrac-
tion called unfused (incomplete) tetanus (tetan- rigid, tense;
Figure 10.15c). When a skeletal muscle fiber is stimulated at a
higher rate of 80 to 100 times per second, it does not relax at all.
The result is fused (complete) tetanus, a sustained contraction
in which individual twitches cannot be detected (Figure 10.15d).
Wave summation and both kinds of tetanus occur when
additional Ca 2 is released from the sarcoplasmic reticulum by
subsequent stimuli while the levels of Ca2 in the sarcoplasm
are still elevated from the first stimulus. Because of the buildup
in the Ca2 level, the peak tension generated during fused
tetanus is 5 to 10 times larger than the peak tension produced
during a single twitch. Even so, smooth, sustained voluntary
muscle contractions are achieved mainly by out-of-synchrony
unfused tetanus in different motor units.
The stretch of elastic components, such as tendons and con-
nective tissues around muscle fibers, also affects wave summa-
tion. During wave summation, elastic components are not given
much time to spring back between contractions, and thus remain
taut. While in this state, the elastic components do not require
very much stretching before the beginning of the next muscular
contraction. The combination of the tautness of the elastic com-
ponents and the partially contracted state of the filaments
enables the force of another contraction to be greater than the
one before.

Figure 10.15 Myograms showing the effects of different frequencies of stimulation. (a) Single twitch. (b) When a second stimulus
occurs before the muscle fiber has relaxed, the second contraction is stronger than the first, a phenomenon called wave
summation. (The dashed line indicates the force of contraction expected in a single twitch.) (c) Unfused tetanus produces
a jagged curve due to partial relaxation of the muscle fiber between stimuli. (d) In fused tetanus, which occurs when
there are 80–100 stimuli per second, the myogram line, like the contraction force, is steady and sustained.
Due to wave summation, the tension produced during a sustained contraction is greater than that produced by a
single twitch.

Myograms
Force of contraction

Action
potential

Time (msec)

(a) Single twitch (b) Wave summation (c) Unfused tetanus (d) Fused tetanus

? Would the peak force of the second contraction in (b) be larger or smaller if the second stimulus were applied a few
milliseconds later?

Motor Unit Recruitment
The process in which the number of active motor units increases
is called motor unit recruitment. Typically, the different motor
units of an entire muscle are not stimulated to contract in unison.
While some motor units are contracting, others are relaxed. This
pattern of motor unit activity delays muscle fatigue and allows
contraction of a whole muscle to be sustained for long periods.
The weakest motor units are recruited first, with progressively
stronger motor units added if the task requires more force.
Recruitment is one factor responsible for producing smooth
movements rather than a series of jerks. As mentioned, the num-
ber of muscle fibers innervated by one motor neuron varies
greatly. Precise movements are brought about by small changes
in muscle contraction. Therefore, the small muscles that produce
precise movements are made up of small motor units. For this
reason, when a motor unit is recruited or turned off, only slight
changes occur in muscle tension. By contrast, large motor units
are active where large tension is needed and precision is less
important.
• CL I N IC AL CO NN EC TI ON Aerobic Training versus
Strength Training
Regular, repeated activities such as jogging or aerobic dancing increase
the supply of oxygen-rich blood available to skeletal muscles for aero-
bic cellular respiration. By contrast, activities such as weight lifting rely
more on anaerobic production of ATP through glycolysis. Such anaero-
bic activities stimulate synthesis of muscle proteins and result, over
time, in increased muscle size (muscle hypertrophy). Athletes who en-
gage in anaerobic training should have a diet that includes an adequate
amount of proteins. This protein intake will allow the body to synthesize
muscle proteins and to increase muscle mass. As a result, aerobic train-
ing builds endurance for prolonged activities; in contrast, anaerobic
training builds muscle strength for short-term feats. Interval training is
a workout regimen that incorporates both types of training—for exam-
ple, alternating sprints with jogging. •
Muscle Tone
Even at rest, a skeletal muscle exhibits muscle tone (tonos
tension), a small amount of tautness or tension in the muscle due
to weak, involuntary contractions of its motor units. Recall that
skeletal muscle contracts only after it is activated by acetyl-
choline released by nerve impulses in its motor neurons. Hence,
muscle tone is established by neurons in the brain and spinal
cord that excite the muscle’s motor neurons. When the motor
neurons serving a skeletal muscle are damaged or cut, the
muscle becomes flaccid (FLAK-sid or FLAS-sid flabby), a
state of limpness in which muscle tone is lost. To sustain muscle
tone, small groups of motor units are alternatively active and in-
active in a constantly shifting pattern. Muscle tone keeps skeletal
muscles firm, but it does not result in a force strong enough to
produce movement. For example, when the muscles in the back
CONTROL OF MUSCLE TENSION 323
of the neck are in normal tonic contraction, they keep the head
upright and prevent it from slumping forward on the chest.
Muscle tone also is important in smooth muscle tissues, such as
those found in the gastrointestinal tract, where the walls of the
digestive organs maintain a steady pressure on their contents.
The tone of smooth muscle fibers in the walls of blood vessels
plays a crucial role in maintaining blood pressure.
• C L I NI C AL CO NN ECT IO N Hypotonia and Hypertonia
Hypotonia (hypo- below) refers to decreased or lost muscle tone.
Such muscles are said to be flaccid. Flaccid muscles are loose and ap-
pear flattened rather than rounded. Certain disorders of the nervous
system and disruptions in the balance of electrolytes (especially
sodium, calcium, and, to a lesser extent, magnesium) may result in flac-
cid paralysis, which is characterized by loss of muscle tone, loss or re-
duction of tendon reflexes, and atrophy (wasting away) and degenera-
tion of muscles.
Hypertonia (hyper- above) refers to increased muscle tone and is
expressed in two ways: spasticity or rigidity. Spasticity (spas-TIS-i-tē) is
characterized by increased muscle tone (stiffness) associated with an
increase in tendon reflexes and pathological reflexes (such as the
Babinski sign, in which the great toe extends with or without fanning of
the other toes in response to stroking the outer margin of the sole).
Certain disorders of the nervous system and electrolyte disturbances
such as those previously noted may result in spastic paralysis, partial
paralysis in which the muscles exhibit spasticity. Rigidity refers to in-
creased muscle tone in which reflexes are not affected, as occurs in
tetanus. •
Isotonic and Isometric Contractions
Muscle contractions are classified as either isotonic or isometric.
In an isotonic contraction (iso- equal; -tonic tension), the
tension (force of contraction) developed by the muscle remains
almost constant while the muscle changes its length. Isotonic
contractions are used for body movements and for moving
objects. The two types of isotonic contractions are concentric
and eccentric. In a concentric isotonic contraction, if the ten-
sion generated is great enough to overcome the resistance of the
object to be moved, the muscle shortens and pulls on another
structure, such as a tendon, to produce movement and to reduce
the angle at a joint. Picking a book up off a table involves con-
centric isotonic contractions of the biceps brachii muscle in the
arm (Figure 10.16a). By contrast, as you lower the book to place
it back on the table, the previously shortened biceps lengthens in
a controlled manner while it continues to contract. When the
length of a muscle increases during a contraction, the contraction
is an eccentric isotonic contraction (Figure 10.16b). During an
eccentric contraction, the tension exerted by the myosin cross-
bridges resists movement of a load (the book, in this case) and
slows the lengthening process. For reasons that are not well un-
derstood, repeated eccentric isotonic contractions (for example,
walking downhill) produce more muscle damage and more de-
324 CHAPTER 10 • MUSCULAR TISSUE

Figure 10.16 Comparison between isotonic (concentric and eccentric) and isometric contractions. Parts (a) and (b) show isotonic
contractions of the biceps brachii muscle in the arm; part (c) shows isometric contraction of shoulder and arm muscles.
In an isotonic contraction, tension remains constant as muscle length decreases or increases; in an isometric
contraction, tension increases greatly without a change in muscle length.

Movement

No Movement

Movement

(a) Concentric contraction while (b) Eccentric contraction while (c) Isometric contraction while
picking up a book lowering a book holding a book steady

? What type of contraction occurs in your neck muscles while you are walking?

layed-onset muscle soreness than do concentric isotonic contrac-

tions.
In an isometric contraction (-metro measure or length),
the tension generated is not enough to exceed the resistance of
the object to be moved, and the muscle does not change its
length. An example would be holding a book steady using an
outstretched arm (Figure 10.16c). These contractions are impor-
tant for maintaining posture and for supporting objects in a fixed
position. Although isometric contractions do not result in body
movement, energy is still expended. The book pulls the arm
downward, stretching the shoulder and arm muscles. The iso-
metric contraction of the shoulder and arm muscles counteracts
the stretch. Isometric contractions are important because they
stabilize some joints as others are moved. Most activities include
both isotonic and isometric contractions.
C H ECK PO INT
15. How are the sizes of motor units related to the degree of
muscular control they allow?
16. What is motor unit recruitment?
17. Why is muscle tone important?
18. Define each of the following terms: concentric isotonic
contraction, eccentric isotonic contraction, and isometric
contraction.
19. Demonstrate an isotonic contraction. How does it feel?
What do you think causes the physical discomfort you
are experiencing?
TYPES OF SKELETAL
MUSCLE FIBERS
OBJECTIVE
• Compare the structure and function of the three types of
skeletal muscle fibers.
Skeletal muscle fibers are not all alike in composition and
function. For example, muscle fibers vary in their content of
myoglobin, the red-colored protein that binds oxygen in muscle
fibers. Skeletal muscle fibers that have a high myoglobin content
are termed red muscle fibers and appear darker (the dark meat
in chicken legs and thighs); those that have a low content of
myoglobin are called white muscle fibers and appear lighter
(the white meat in chicken breasts). Red muscle fibers also
contain more mitochondria and are supplied by more blood
capillaries.
Skeletal muscle fibers also contract and relax at different
speeds, and vary in which metabolic reactions they use to gener-
ate ATP and in how quickly they fatigue. For example, a fiber is
categorized as either slow or fast depending on how rapidly
the ATPase in its myosin heads hydrolyzes ATP. Based on all
these structural and functional characteristics, skeletal muscle
fibers are classified into three main types: (1) slow oxida-
tive fibers, (2) fast oxidative-glycolytic fibers, and (3) fast gly-
colytic fibers.