3.

4 Skeletal Muscle Mechanics

cle actually lengthens. These are called eccentric

Learning Objectives contractions.

G Contrast skeletal, cardiac, and smooth muscles MUSCLES PERFORM DIVERSE

G Describe an isometric contraction
G Describe a muscle twitch
FUNCTIONS
G Distinguish between slow-twitch and fast-twitch muscle Muscles that are connected to the skeleton are called
fibers on the basis of their contractile behavior skeletal muscles. Because they are connected to the
G Define the motor unit bones via their tendons, these muscles transfer force
G Describe how recruitment increases muscle force to the skeleton and move one bone relative to another.
G Describe how a muscle responds to increased frequency of
Coordinated activation of muscles allows for the various
stimulation
movements of the joints (flexion and extension; abduc-
G
tion and adduction; protraction and retraction; elevation
Define tetany
and depression; pronation and supination; medial
G Describe the size principle of motor unit recruitment
rotation and lateral rotation; and circumduction).
G Describe the relative importance of frequency coding ver- Skeletal muscles allow us to lift weights and to move
sus population coding for grading muscle force ourselves from place to place. These muscles also
G Define the innervation ratio and explain how the size prin- include the tongue, the muscles that move the
ciple creates proportional control of muscle force eyeballs, and the upper third of the esophagus. A sec-
G Define passive and active tension and show how they vary ondary function of these skeletal muscles is the
with muscle length production of body heat. During vigorous exercise,
G Define afterload the heat produced by the skeletal muscles raises body
G Describe an isotonic contraction temperature and the excess heat must be removed
G Draw the force
velocity curve for fast-twitch and slow- from the body.
twitch muscles Other muscles surround hollow organs. These include
G Show how power varies with velocity of contraction the heart, urinary bladder, gallbladder, and uterus.
G Distinguish among concentric, isometric, and eccentric Contraction of the muscles in the walls of these organs
contractions develops a tension in the walls and so develops pres-
G Describe muscle pinnation and how this contributes to sure within the organ. Still other muscles are present
muscle function in the long hollow tubes of the body. Contraction of
G Define muscle fatigue the muscles in these tubes sets the diameter of a tube
G Describe Burke’s system of muscle classification based on and thereby controls flow through it. Examples include
contractile properties the muscular sphincters of the gastrointestinal tract,
the arterioles, the ureters, and the airways of the lungs.
Contraction of the muscular layers of the intestine also
propels material through the intestine.
MUSCLES EITHER SHORTEN OR
PRODUCE FORCE MUSCLES ARE CLASSIFIED ACCORDING
The primary action of a muscle is to contract. Usually TO FINE STRUCTURE, NEURAL
this word means to shorten, but physiologists often use CONTROL, AND ANATOMICAL
it in a broader sense, meaning to activate the muscle. ARRANGEMENT
Activation of the muscles can produce force without
actually shortening, as you might do when holding STRIATED MUSCLES HAVE STRIPES
something really heavy. As we shall see later, there is a Under the microscope, some muscles appear to have
trade-off between shortening and producing force, and stripes running across them. These are said to be stri-
both originate from the same mechanism. In everyday ated (from the Latin meaning channel or furrow). All of
motion, some muscle contractions occur while the mus- the skeletal muscle and cardiac muscle are striated.
292
© 2017 Elsevier Inc. All rights reserved.
DOI: http://dx.doi.org/10.1016/B978-0-12-800883-6.00027-6
 S k e l e t a l M u sc l e M e c ha ni c s 293
Muscles lacking stripes are called smooth muscles ISOMETRIC FORCE IS MEASURED
(Figure 3.4.1).
WHILE KEEPING MUSCLE LENGTH
SKELETAL MUSCLES ARE USUALLY ACTIVATED CONSTANT
VOLUNTARILY A muscle can be dissected free from its origin and inser-
Most skeletal muscles are normally activated by a motor tion (the places where the tendons attach to the bones)
neuron that resides in the anterior or ventral horn of and connected to a device to measure the force devel-
the spinal cord. Commands for the activation of the oped upon activation, as shown in Figure 3.4.2. Here
motor neuron can come from sensory input, as in the the experimental setup measures force at a nearly con-
spinal reflexes, or from the motor cortex in the cere- stant length, and so these are called isometric contrac-
brum. These muscles are organized as motor units, tions (from “iso,” meaning “the same,” and “metric”
with nerve fibers providing the signal for activation. In meaning “distance”). When a stimulus is turned on very
contrast, cardiac and visceral smooth muscles are intrin- briefly (3 ms), a single muscle contraction, called the
sically active and do not require neural initiation of acti- muscle twitch, is recorded. Force rises and falls without
vation. However, these muscles are innervated and their muscle shortening. It takes some time for force to rise
activity is modulated by nerves and sometimes by and generally a longer time for it to fall back to resting
humoral factors. These different types of control are levels. When a gastrocnemius muscle of a rat is used,
broadly classed as voluntary and involuntary. the twitch time is about 70 ms. When the soleus is
Breathing and blinking the eyelids are largely automatic used, the twitch time is over 150 ms. Muscles can be
events that occur involuntarily, but we can consciously distinguished on the basis of their contractile properties,
suppress these activities for some time and we must be including their twitch times. The gastrocnemius is an
able to control breathing voluntarily in order to speak, example of a fast-twitch muscle; the soleus is a slow-
sing, or swim. This chapter is devoted to voluntary skel- twitch muscle.
etal muscles. The recording in Figure 3.4.2 also shows that the action
potential on the muscle lasts only a few ms, whereas
the twitch lasts much longer. The action potential is a
Classification of muscle trigger for the muscle contraction.
Neural control Anatomical Histological
MUSCLE FORCE DEPENDS ON THE
NUMBER OF MOTOR UNITS THAT ARE
Voluntary Skeletal
ACTIVATED
Striated
Cardiac
The stimulus applied to the nerve can be varied by
increasing its voltage or changing its duration, delay, or
Involuntary
frequency. When the voltage strength of the stimulus is
Visceral Smooth
gradually increased, the force of the twitch also
FIGURE 3.4.1 Classification of muscles can be based on control increases until the force reaches a plateau, as shown in
properties, anatomical properties, or the fine structure observed Figure 3.4.3. Why should muscle force depend on the
histologically. The preferred classification is that highlighted in the voltage of the stimulus?
figure, which is a mixture of several classification schemes.

Stimulator
+
-

Fast-twitch muscle
Skeletal muscle
Nerve Organ bath Slow-twitch muscle
AP
Force transducer

Recorder

0 50 100 150 200

Time (ms)

FIGURE 3.4.2 Experimental setup for measuring isometric tension in an isolated muscle. In this preparation, the nerve that usually activates the muscle
is severed and action potentials on the motor nerve axon must be initiated by an external stimulator. The muscle is tied at one end to a rigid support
and at the other end to a force transducer. When the muscle contracts, it pulls against a stiff spring located in the transducer. The transducer turns its
slight movement into an electrical signal that can be recorded. Because the actual shortening of the spring, and its attached muscle, is extremely
small, this contraction is called an isometric contraction. A single stimulation results in an action potential in the nerve and subsequently in the muscle
that activates the muscle to produce a single contraction called a twitch. Different muscles differ markedly in the time course of force development
and relaxation.
294 QUANTITATIVE HUMAN PHYSIOLOGY: AN INTRODUCTION

Volts of stimulation
1 2 3 4 5 6 8 10 15 2030 40

Twitch
Vary voltage force
Stimulator
+
-
100 g
Skeletal muscle
Nerve Organ bath
Force transducer
Time (s)

600
Recorder 500
400

Force (g)
300
200
100
0
0 10 20 30 40
Stimulation intensity(v)

FIGURE 3.4.3 Increase in the muscle twitch with increased recruitment of motor fibers by increasing the strength of the external stimulus. Data
obtained from electrical stimulation of the sciatic nerve serving the gastrocnemius muscle of the rat.

Increasing strength of stimulus

progressively activates axons
further from the electrode until all
axons are finally activated At any given stimulus strength,
small axons reach threshold more easily
because they are more excitable

Extracellular electrode
Epineurium
Perineurium

Endoneurium

Myelinated axons

FIGURE 3.4.4 Intact nerve and the extracellular electrode that stimulates it. The whole nerve is a bundle of axons originating in either sensory or
motor neurons. As the strength of stimulus increases, more axons are activated until, eventually, all axons in the nerve fire action potentials. Since
each axon innervates a set of muscle fibers, called its motor unit, progressive activation of axons causes progressive increases in the number of
activated muscle fibers, and progressive increases in the total force produced by the muscle. The smaller axons are more excitable, and they are
activated first.

The motor nerve supplying the muscle is actually a bun- across their membranes. But the current from the extra-
dle of axons from motor neurons carrying excitation to cellular electrode has many places to go, and most of
the muscle and sensory nerve fibers that return informa- the current does not go across the axon membrane.
tion to the central nervous system about the state of the Thus higher voltages are necessary to depolarize all of
muscle. The proportion of motor nerve axons within the axons in the bundle. This is why volts are necessary
the bundle varies from 40% to 70% in different mus- for the external stimulus, when only a few millivolts of
cles, with the remaining being axons carrying sensory membrane depolarization are sufficient to initiate an
information. The motor axons come in two general action potential. Further, the different sized motor neu-
types: small and large diameter axons (see Figure 3.4.4). ron axons differ in their rheobase—the amount of cur-
The stimulus that we apply is an extracellular stimulus rent for an infinite time that will bring the axons to
that depolarizes the axons by passing an inward current threshold (see Figure 3.2.8). Increasing the stimulus
 S k e l e t a l M u sc l e M e c ha ni c s 295

Motor neuron 1 Motor neuron 2

A
B

D
C

Muscle fibers with striations

FIGURE 3.4.5 The motor unit consists of the motor neuron and all of the muscle fibers innervated by it. Here motor neuron 1 innervates fibers A, B,
and C; motor neuron 2 innervates fibers D and E. When only motor neuron 1 fires an action potential, only fibers A, B, and C contribute to the force.
When only motor neuron 2 fires an action potential, only fibers D and E contribute to the force developed by the muscle. When both motor neurons
fire an action potential, all of the muscle fibers contribute to the force. Thus the force is greater when more motor units are recruited.

strength (voltage) increases the number of motor neu- that are activated depends on the output of the primary
rons that are activated, which increases the number of motor cortex, along with modulating influences from
muscle fibers that are activated, which increases force. other brain areas and from sensory information. The
Each motor nerve branches and connects to a set of neural control of movement is discussed in Chapter 4.5.
muscle fibers, making a neuromuscular junction with
each of them. The motor neuron and the set of muscle
fibers it innervates make up a motor unit. The entire THE SIZE PRINCIPLE STATES THAT
muscle consists of a large number of muscle fibers that MOTOR UNITS ARE RECRUITED IN THE
are each typically innervated by a single motor neuron ORDER OF THEIR SIZE
making a junction in the middle of the muscle fiber.
Motor neurons typically innervate more than one mus- Large motor units are innervated by large motor neu-
cle fiber, but each muscle fiber is innervated by only rons, and smaller motor units are innervated by smaller
one motor neuron. The motor unit is designed for motor neurons. The small motor neurons are more
100% fidelity: each time the motor neuron is activated, excitable, so these are recruited first. This corresponds to
all of its muscle fibers are subsequently activated. Thus our everyday experience. When trying to perform deli-
motor units are indivisible quantal elements in all cate movements that require dexterity but little force,
movements (see Figures 3.4.4 and 3.4.5). control of muscle force must be fine. This is accom-
plished by recruiting small numbers of muscle fibers.
The increase in force by activating increasing number of When performing gross motor movements involving a
motor units is called recruitment. The nearly continu- lot of force, the increments of force are large and we
ous variation in force from small to large forces means recruit successively larger motor units. The recruitment
that this force is not all or none; it is graded. Although of motor units in order of their sizes is accomplished
the gradation is fine, it is not really continuous because through other nerves that make connections to the
muscle fibers are activated in a discrete (as opposed to lower motor neurons.
continuous) way. Because there are so many muscle
fibers, the force appears to vary nearly continuously. It
is important to note that the method of recruitment MUSCLE FORCE CAN BE GRADED BY
here, increasing the voltage of stimulation of the nerve, THE FREQUENCY OF MOTOR NEURON
is not physiological. Ordinarily motor units are recruited
through neuronal connections to the motor neurons,
FIRING
particularly by the activation of command signals origi- The action potential on the motor neuron is very
nating from the primary motor cortex in the brain, short, between 1 and 3 ms. The action potential on the
located in the precentral gyrus (see Chapter 4.5). This muscle cell membrane is also short, on the order of
area of the brain contains “upper” motor neurons that 3
5 ms. The muscle twitches are long by comparison,
provided excitation to the “lower” motor neurons that some 30
300 ms. This means that it is possible to stim-
are located in the ventral horn of the spinal cord and ulate the muscle with another action potential before
directly activate muscle fibers. The motor unit specifi- the muscle has relaxed. Indeed, we can stimulate a
cally refers to a lower motor neuron and the set of mus- muscle again before it reaches its peak tension. When
cle fibers that it innervates. Which lower motor neurons a muscle is stimulated before it has completely relaxed,
296 QUANTITATIVE HUMAN PHYSIOLOGY: AN INTRODUCTION

5 Hz 10 Hz 15 Hz 20 Hz30 Hz 40 Hz 50 Hz 60 Hz
Maximum voltage, 100 g
Stimulator vary frequency Muscle
+ force
- Skeletal muscle
Nerve Organ bath
Force transducer

700
Recorder
600
500

Force (g)
400
300
200
100
0
0 10 20 30 40 50 60
Frequency (Hz)

FIGURE 3.4.6 Gradation of muscle force by the frequency of stimulation. Increasing the frequency of stimulation at low frequencies does not increase
the force; it merely increases the frequency of the same twitch wave form. When the period of the stimulation frequency is shorter than the period of
the twitch, however, force begins to summate. With continued increases in frequency, there is added force until tetany is reached. Data obtained from
electrical stimulation of the sciatic nerve serving intact gastrocnemius muscle of the rat.

a new twitch begins force where the first twitch left As you can see from Figure 3.4.6, the tetanic force is
off (see Figure 3.4.6). The resulting force is greater much greater than the twitch force. In general, tetanic
than with a single twitch. Thus muscle force summates force is about five times the twitch force, but the
with repetitive stimulation. When the frequency of stim- twitch ratio varies from 2 to 10 in different muscle
ulation is great enough, the muscle produces a single types.
forceful contraction with no waviness in the force.
This condition is called tetany.
The frequency required to reach tetany depends on how MUSCLE FORCE DEPENDS ON THE
fast the muscle contracts and relaxes. If the muscle LENGTH OF THE MUSCLE
twitch is 100 ms long, then summation will just begin
The device shown in Figures 3.4.2, 3.4.3, and 3.4.6 can
when the next stimulation arrives at the end of relaxa-
be adjusted to vary the length of the muscle. When
tion. Since there are 10 100 ms-intervals in a second,
muscles are relaxed, they exert no force. When the
summation for such a fiber should begin at a stimula-
relaxed muscles are stretched passively, without activa-
tion frequency of about 10 Hz. (1 Hz is hertz, meaning
tion by a nerve stimulus, they produce a passive force.
a cycle per second; 10 Hz is therefore 10 cycles or events
This is due to elastic properties of the muscle material
per second.) Typically most muscles in the human teta-
itself. This passive force increases steeply and nonli-
nize between 20 and 100 Hz. Summation begins when
nearly with increases in length (see Figure 3.4.7).
stimulation frequency just exceeds the inverse of the
twitch time. When a stretched muscle is stimulated tetanically, it
produces a force in addition to the passive force. This
Although the results shown in Figure 3.4.6 are a labora-
increment in force is called the active tension because it
tory observation involving an isolated nerve and its
depends on activation of the muscle by the tetanic stim-
muscle, the same phenomenon also occurs in living,
ulation. The additional active force produced by stimu-
breathing people. We can grade the force of a muscular
lation depends on the length of the muscle. The active
contraction, or vary force more or less continuously, by
force increases to a maximum and then declines with
altering the frequency of motor neuron firing. This is
further increases in muscle length. The relationship
one of the principal ways of physiological control of
between active force and muscle length is the
muscle force. Since action potentials are an all-or-none
length
tension curve, originally determined in isolated
phenomenon, information can be coded only through
muscle fibers by Robert Ramsey and Sybil Street. This
the frequency of action potentials (frequency code) and
data became an important observation for explaining
in the population of neurons carrying it (population
the mechanism by which muscles produce force.
code—this is the same as recruitment). In the case of
muscle, high-frequency neuronal activity is converted to The resting length of a muscle is usually designated as
high intensity of muscle force. L0. Because muscles are attached to the skeleton, their
 S k e l e t a l M u sc l e M e c ha ni c s 297

Total tension

2500
Stimulator Use maximum voltage, tetanic frequency
2000

+
Active tension

-
Skeletal muscle

Force (g)
1500
Nerve Organ bath
Force transducer 1000
Passive tension
500

0
Recorder 0 2 4 6 8 10 12 14 16
Normalized length (mm)

Vary length
Active tension is the increment
in force upon stimulation
37
36
Muscle
35
force
34
250 g = 2.45 N 33
32
30 31
28 29 Baseline
22 23 24 25 26 27
Passive tension increases
with stretch

FIGURE 3.4.7 The length
tension relationship in muscle. Increasing muscle length from slack length to relatively long lengths causes a passive force
or tension that does not require activation of the muscle. Activation of the muscle at maximum recruitment and tetanic frequency (so that recruitment
or frequency response does not confound the results) causes an increment in the force called the active force or tension. Muscle length was increased
in increments of 1 mm as indicated by the Vernier scale readings under each tetanic stimulation. Active tension increases biphasically with muscle
length. Passive tension increases progressively with muscle length—muscles do not obey Hooke’s law, in which force is proportional to length. The
total tension is the sum of the passive and active forces. Data obtained by electrical stimulation of the sciatic nerve innervating the gastrocnemius
muscle of the rat.

degree of shortening is defined by the movement of the Chapter 3.7) interrupted by rest periods. These series of
bones and the attachment points of the muscles. Most activations increase force by wave summation. Although
muscles do not shorten or lengthen by more than about this is important, it provides a range of forces that scale
25% of their rest length. Thus although muscle length according to the tetanus
twitch ratio, which varies
can determine muscle force, typically the muscle is from about 2
10, depending on the muscle. Therefore,
physiologically arranged near the top of the length
ten- the frequency of activation provides at most about a 10-
sion curve, and muscle length changes are relatively fold range of variation in muscle force. Skeletal muscles
unimportant compared to recruitment and frequency of often have a range of forces that varies 20,000 fold,
stimulation. from the weakest contraction to the most forceful. Thus
rate coding or frequency coding provides an important
part of muscle force gradation, but population coding
RECRUITMENT PROVIDES THE (recruitment) provides the greater part of muscle force
GREATEST GRADATION OF MUSCLE gradation.
FORCE
We have described three distinct ways to vary muscle MUSCLE FIBERS DIFFER IN
force: (1) recruit muscle fibers by activating larger num- CONTRACTILE, METABOLIC AND
bers of motor units, (2) vary the frequency of activation
of the activated motor units, and (3) vary the length. As
PROTEOMIC CHARACTERISTICS
mentioned above, muscle length typically does not vary As shown in Figure 3.4.2, muscles differ in their con-
significantly, and the resting length of muscles is situ- tractile properties. This mechanical difference derives
ated at the top of the active length
tension curve, so from differences in their constituent motor units. Each
that varying muscle length is not an important way of motor neuron innervates a number of muscle fibers.
varying muscle force in the whole person. Most activa- The muscle fibers themselves are heterogeneous, and
tion of muscle is not by a single action potential to pro- several classification schemes have been devised to
duce a twitch, but by a train of impulses (see describe them. Based on contractile behavior, muscle
298 QUANTITATIVE HUMAN PHYSIOLOGY: AN INTRODUCTION

fibers can be broadly classified as being S, FR, FI and 140

FF: these mean S 5 slow; FR 5 fast, fatigue resistant;
FI 5 fast, intermediate in fatigability; FF 5 fast, fatigable. 120
The differences in the muscle fibers come from their
expression of different protein isoforms or from differ- 100

Total force (N)

ing quantities of the same proteins or from differing
80
quantities and composition of subcellular organelles.
The muscle fibers differ in metabolic ability that shows 60
up in their gross appearance: red muscle contains a lot
of myoglobin, a protein that binds oxygen within the 40
cytoplasm of the muscle fiber and aids in oxygen deliv-
ery to mitochondria, which are also rich in red muscles. 20
White muscle fibers rely less on oxidative metabolism
and use anaerobic pathways more. So muscles can be 0
0 20 40 60 80 100
classified according to their metabolic properties as
Percent motor units recruited
being SO 5 slow, oxidative; FOG 5 fast, oxidative and
glycolytic; and FG 5 fast, glycolytic. Muscles are also dif- FIGURE 3.4.8 Simulated active force of a muscle as a fraction of motor
ferentiated on the basis of myosin isoform expression as units that are activated. The muscle was simulated as a population of
type I (equivalent to SO), type IIa (equivalent to FOG), 100 motor units whose aggregate muscle fiber area was given as
Ai11 5 1.05 Ai, with a constant specific force of 20 N cm22.
and type IIb (FG). We will return to these different mus-
cle fiber types in Chapters 3.5 and 3.7.

their muscle fibers have a smaller cross-sectional area.

MOTOR UNITS CONTAIN A SINGLE The first recruited motor units typically have Type I
TYPE OF MUSCLE FIBER muscle fibers (slow twitch, oxidative metabolism) and
Muscle fibers are heterogeneous, differing in contractile the last recruited motor units innervate large muscle
and metabolic properties, but all of those muscle fibers fibers of Type IIB (fast, glycolytic metabolism).
that are innervated by a single motor neuron are of a Innervation ratios vary with the type of muscle from
single type. This fact suggests that there is some kind of about 5 in the lateral rectus muscle that moves the eye-
communication that passes between the motor neuron ball to over 2000 muscle fibers for a single motor unit
and the muscle fibers that directs their expression of in the medial gastrocnemius in the calf. According to
protein isoforms. In short, the motor neuron establishes the size principle, the small motor units are recruited
the kind of muscle fibers it innervates and maintains first, and progressively larger motor units are recruited
them in that form. until all motor units are recruited in maximal voluntary
contractions. The theoretical consequence of the disper-
sion of motor unit innervation ratios and the orderly
THE INNERVATION RATIO OF MOTOR recruitment of motor units is shown in Figure 3.4.8.
UNITS PRODUCES A PROPORTIONAL The total active force produced by the muscle increases
CONTROL OF MUSCLE FORCE progressively with the recruitment of the motor units in
a process referred to as proportional control. In this
The tetanic force produced by a single motor unit is process, force is added in steps that are proportional to
given as the amount of force already present. During weak con-
½3:4:1 FMU 5 NFMF 5 NAFS tractions, force is adjusted by changing it slightly (e.g.,
in steps of 5%) whereas strong contractions are adjusted
where FMU is the force of the motor unit, N is the num- by making large incremental changes that remain 5% of
ber of muscle fibers that the motor neuron innervates, the force already present. The proportional control is
A is the area per muscle fiber, and FS is the specific force accomplished by recruiting motor units in order of
or the force per unit area of muscle fiber. N is the increasing strength.
innervation ratio. Although some reports suggest that
there might be differences in the specific force for differ-
ent muscle fiber types, the effect is not large. Typically MUSCLE FORCE VARIES INVERSELY
the force per unit area for a muscle fiber is about
20 N cm22. The total force developed by a muscle is WITH MUSCLE VELOCITY
given as The experimental setups illustrated in Figures 3.4.2,
3.4.3, 3.4.6, and 3.4.7 were all performed using isomet-
X
i5j
ric contractions in which muscle length did not change.
½3:4:2 F5 Ni Ai FS
We now change this situation and hook up one end of
i51
the muscle to a pulley system connected to an afterload
indicating that the forces of the motor units from unit 1 (see Figure 3.4.9). An afterloaded muscle cannot
to j sum linearly, and that the innervation ratio and shorten until it produces a force equal to the afterload.
area of the muscle fibers vary with motor unit. Typically When stimulated, the muscle contracts isometrically
the small motor units have low innervation ratios and during the time it takes to develop force equal to the
 S k e l e t a l M u sc l e M e c ha ni c s 299

Velocity of shortening
is the slope of L vs t

Time (s)
0 0.5 1.0 1.5 2.0 2.5 3.0

Length
Position, X
As afterload increases,
Use maximum voltage, tetanic frequency Stimulus isometric phase is longer
Stimulator Stimulus Stimulus
and isotonic force is higher
+
-

Isotonic phase
Isometric phase
Organ bath

Isometric phase
Force transducer

isometric phase
Nerve

Isotonic phase
Isotonic phase
Force

Afterload skeletal muscle Recorder

0 0.5 1.0 1.5 2.0 2.5 3.0

Time (s)

FIGURE 3.4.9 Experimental setup for measuring the force and velocity of isotonic contractions. The afterload is supported by a shelf prior to the
activation of the muscle and thus it is not felt by the muscle until all slack in the line are taken up by muscle shortening. When the size of the
afterload is increased, the muscle takes longer to develop sufficient force to lift the load, and the velocity of shortening is less. The initial velocity of
shortening is inversely related to the afterload or the force developed by the muscle.

afterload. After the muscle develops a force equal to the

afterload, it lifts the afterload and continues to shorten. Maximum velocity occurs at no load
For some part of this contraction, the velocity of short- 50
ening is approximately constant. Such a contraction is
Velocity (cm s–1)

called an isotonic contraction. Thus each contraction 40

consists of an isometric part and an isotonic part, as
30 Fast-twitch fibers Maximum force occurs
shown in Figure 3.4.9.
at zero velocity
Everyday experience shows that the speed of muscle 20
contraction depends on the load that must be moved.
We know that we can move a light load quickly, 10 Slow-twitch
whereas we move a heavy load slowly. The force
velo-
city curve is produced by measuring the initial velocity 0
and plotting it against the afterload. The initial velocity 0 10 20 30 40
can be measured using a device such as that shown in Force (N cm–2)
Figure 3.4.9, where the afterload is varied. The initial FIGURE 3.4.10 The force
velocity curve for an intact fast-twitch or
velocity is used because after the muscle shortens it can- slow-twitch muscle. Fast contractions can occur only with low loads.
not produce as much force, because it moves off the Higher loads mean slower velocity of muscle contraction, but more
optimum of the length
tension curve. The force
velo- force. Thus there is an inverse relationship between force and velocity.
city curve is shown in Figure 3.4.10.
Inserting Eqn [3.4.3C] into Eqn [3.4.3B] and then into
MUSCLE POWER VARIES WITH THE Eqn [3.4.3A], we get
LOAD AND MUSCLE TYPE ½3:4:4 Power 5 force 3 distance=time
Power is defined as the rate of energy production or con- From the definition of velocity as distance/time, we get
sumption and it has the units of energy per unit time.
In mechanical terms, energy is work. Work is defined in ½3:4:5 Power 5 force 3 velocity
mechanics as force 3 distance. Thus we have the
following: The force
velocity curve obtained experimentally is dis-
played in Figure 3.4.10. We can obtain the power
force
½3:4:3A Power 5 Energy=time curve by multiplying force and velocity at every point
on this curve. This power is the instantaneous power
½3:4:3B Energy 5 work produced during the initial shortening of the muscle.
½3:4:3C Work 5 force 3 distance Power is expressed in units of watts or N m s21, per
unit weight of muscle. The power as a function of
muscle force is shown in Figure 3.4.11. This shows
300 QUANTITATIVE HUMAN PHYSIOLOGY: AN INTRODUCTION

that the power is about two to three times greater in

Maximum power peaks at about
one-third maximum force
fast-twitch fibers because of their greater speed of
contraction.
Fast-twitch muscle 400
50 ECCENTRIC CONTRACTIONS LENGTHEN
300 THE MUSCLE AND PRODUCE MORE
Velocity (cm s–1)

40
FORCE

Power (W kg–1)
30 According to the way in which we measured velocity,
200
Slow-twitch a positive velocity corresponded to a shortening of
20
the muscle. If the support for a very large afterload in
100 Figure 3.4.9 is removed, the afterload will cause the
10
muscle to lengthen during contraction. This length-
0 0 ening is a negative velocity. Contraction of the mus-
0 10 20 30 40 cle during a lengthening is called an eccentric
Force (N cm–2 ) contraction. Contraction of a muscle that causes a
shortening is called a concentric contraction. The
FIGURE 3.4.11 Power as a function of force for slow-twitch and fast- extension of the force
velocity curve to negative
twitch muscles. Velocity is shown as dashes, power as solid lines. Power
peaks at about one-third maximum force for both fast-twitch and slow- velocities (see Figure 3.4.12) shows that muscles can
twitch muscles, but fast-twitch muscles deliver more power due to their exert about 40% more force in an eccentric contrac-
faster contractions. These curves are part of the reasons for gears on tion compared to the maximal isometric force
bicycles, so that a constant power can be delivered to the wheels by measured at zero velocity.
keeping the velocity of the muscles and forces near the peak power
output when translational velocity of the bike changes going up or
down hill. CONCENTRIC, ISOMETRIC, AND
ECCENTRIC CONTRACTIONS SERVE
DIFFERENT FUNCTIONS
Because concentric contractions shorten, they are use-
Velocity (cm s–1)

50 ful for the acceleration of one body part relative to

another, including parts that are loaded with external
40 objects. Isometric contractions are used to fix joints,
usually to produce a platform on which other actions
30 Concentric contraction can be made. For example, delicate work by the fin-
gers requires immobilization of the arm and shoulder
Shortening

20
Isometric contraction
to hold the hand still while the fingers do the work.
10 Such immobilization is accomplished by simulta-
neously activating antagonistic muscles—those that
0
move joints in opposite directions. Eccentric contrac-
Lengthening

10 20 30 40
–10 Force tions are used to decelerate body parts, as in activa-
(N cm–2) tion of the quadriceps muscles in the leg while going
–20 Eccentric contraction downstairs.
Table 3.4.1 shows the three types of contractions, their
FIGURE 3.4.12 Concentric, isometric, and eccentric contractions. functions for movement, and the work performed.
Concentric contractions involve a shortening of the muscle. Eccentric
contractions involve a lengthening of the muscle. Isometric contractions
occur when the muscle length does not change and occurs at zero MUSCLE ARCHITECTURE INFLUENCES
velocity. Developed force is greatest for eccentric contractions, next FORCE AND VELOCITY OF THE WHOLE
highest for isometric contractions and lowest for concentric contractions.
MUSCLE
The force that a muscle develops depends on its size:
larger muscles produce greater force. Because muscles

TABLE 3.4.1 Types of Contractions and Their Uses

Types of Contractions Distance Change Function Work

Concentric Shortening (1D) Acceleration Positive W 5 F 3 (1D)
Isometric No change (0 D) Fixation Zero
Eccentric Lengthening (2D) Deceleration Negative W 5 F 3 (2D)
 S k e l e t a l M u sc l e M e c ha ni c s 301
are not right cylinders but take on complicated 300 cm3 and a resting length of 36.8 cm and a second
shapes, determinants of the size present something of muscle with pinnate architecture with fibers 12 cm
a problem. The usual estimate of size is the muscle’s long and overall length of 36.8 cm. The pinnate mus-
cross-sectional area at the belly, or widest part, of the cle fibers are oriented 15 to the line of action. The
muscle. The maximum force expressed per unit area geometry of these is shown in Figures 3.4.14 and
is typically from 15 to 40 N cm22. This is enough to 3.4.15. The fibers in the strap muscle do not span the
lift about 1.5
4 kg cm22. The variation in this force distance from tendon to tendon. Instead, long strap
per unit area derives from the orientation of the muscles are divided into compartments by fibrous
muscle fibers within the muscle. bands called inscriptions. The sartorius muscle (origi-
nating on the lateral hip, wrapping around to the
There are three major orientations of the muscle fibers
inner thigh, and inserting on the medial tibia) has
within muscles: parallel fibers, fusiform, and pin-
three inscriptions, giving four compartments; the
nate. The parallel arrangement is present in muscles
semitendinosus (part of the hamstring muscles, origi-
shaped like a strap or in parts of flat-shaped muscles.
nating on the medial tuberosity of the ischium and
In fusiform muscles, the fibers are generally parallel to
inserting its long tendon on the medial tibia) has
the longitudinal axis of the muscle, but their number
three compartments and the biceps femoris (in the
varies with distance from the belly of the muscle. In
back of the thigh, originating in two places and insert-
pinnate muscles, the fibers are oriented at an angle to
ing on the lateral side of the head of the fibula) and
the tendon or aponeurosis. Because they resemble a
gracilis (most superficial muscle on the inner thigh,
feather, these muscles are called pinnate or pennate
originating on the symphysis pubis and inserting on
(see Figure 3.4.13).
the medial tibia) have two compartments each.
The consequence of orienting the muscle fibers at an
The conclusion from the calculations in Examples 3.4.1
angle with respect to the tendon is to increase the effec-
and 3.4.2 is that the pinnate arrangement of the muscle
tive cross-sectional area of the muscle while reducing
fibers allows many more muscle fibers to attach to the
the distance the muscle can contract along the lines
aponeurosis. The consequence is that the muscle can
of the tendons. To see this, we consider two muscles, a
develop more force, but it does so by reducing its velocity.
strap muscle with parallel fibers having a volume of

Tendons

Bipennate

Muscle fibers
FIGURE 3.4.13 Different arrangement of muscle fibers. Parallel
Pinnate fibers are oriented longitudinally in the direction of the muscle.
Fusiform muscles are tapered. Pinnate fibers are parallel but
Parallel fibered Fusiform oriented at an angle to the action of the muscle.

L = 24.8 cm
L = 36.8 cm Area = 12.1 cm2
Area = 8.15 cm2

Contracted
Rest
FIGURE 3.4.14 Geometry of a strap
muscle with two inscriptions, fiber
Lfiber = 12 cm Lfiber = 8 cm
length 5 12 cm and volume 5 300 cm2.
302 QUANTITATIVE HUMAN PHYSIOLOGY: AN INTRODUCTION

(A) Area = 96.6 cm2

Rest Gross muscle

L = 36.8 cm Cross-sectional area = 11.9 cm2

Lfiber = 12 cm
12 sin 15° = 3.11 cm
15°

36.8–11.59 = 25.21 cm 12 cos 15° = 11.59 cm

Total cross-sectional area = Area x sin 15° = 96.6 cm2 x sin15° = 25 cm2

(B) Area = 101 cm2

Contracted
L = 33.2 cm

Lfiber = 8 cm
22°
FIGURE 3.4.15 Geometry of a pinnate muscle Gross muscle
fiber at rest (A) and contracted (B). CSA = 9.0 cm2

EXAMPLE 3.4.1 Calculate the Isometric Force and Maximal Velocity of a Parallel Muscle
We use the strap muscle shown in Figure 3.4.14 as an example. F 5 8:15 cm2 3 20 Ncm22 5 163 N
Its maximal force is its cross-sectional area times the force devel-
oped per unit cross-sectional area. Since its volume is 300 cm3 Taking the unloaded contraction time as 0.033 s for the muscle
and its length is 36.8 cm, its cross-sectional area is 300 cm3/ fibers to contract from 12 to 8 cm, we get an unloaded
36.8 cm 5 8.15 cm2. velocity 5 (36.8 cm
24.8 cm)/0.033 s 5 364 cm s21.

The typical isometric force per unit area is 20 N cm22. The iso-
metric force of this muscle would be

EXAMPLE 3.4.2 Calculate the Isometric Force and Maximal Velocity of a Unipinnate Muscle
An unipinnate muscle with the same volume and rest length as is the cross-sectional area of the fibers 3 the force per unit
the strap muscle has fibers oriented at 15 to the direction of area 5 25 cm2 3 20 N cm22 5 500 N. This force is not directed
force. The fiber rest length is 12 cm; thus the aggregate cross- along the lines of the aponeurosis; the force transmitted to the
sectional area of the fibers is 300 cm3/12 cm 5 25 cm2, which can tendons will be 500 N 3 cos 15 5 483 N.
also be obtained from the area of the aponeurosis 3 sin 15 . This
The apparent isometric force of the muscle, using its gross cross-
is not the gross cross-sectional area of the muscle. The gross cross-
sectional area, is 483 N/11.9 cm2 5 40.6 N cm22.
sectional area can be estimated from the volume and length of
the muscle. From the geometry in Figure 3.4.15, the overall length When the fibers shorten from 12 to 8 cm, the pinnation angle
of the muscle is 36.8 cm, but each aponeurosis is 25.21 cm long; changes. The overall length of the muscle changes from 36.8 to
the thickness of the muscle is 3.11 cm. From these numbers, the 33.2 cm, or by 3.6 cm, in 0.033 s, giving an unloaded velocity of
area of the aponeuroses is 96.6 cm2 and the gross cross-sectional 3.6 cm/0.033 s 5 109 cm s21.
area of the muscle is 11.9 cm2. Total force generated by the fibers
 S k e l e t a l M u sc l e M e c ha ni c s 303

MUSCLES DECREASE FORCE UPON more motor units; (2) by increasing the frequency of
stimulation; and (3) by changing the length of the mus-
REPEATED STIMULATION; THIS IS cle. Changing the length of the muscle is not so impor-
FATIGUE tant because muscles are restrained by the position of
Everyday experience shows us that maximal effort can their origins and insertions on the skeleton. Recruitment
be sustained only briefly. The more intense the effort, follows the size principle: small motor units are recruited
the faster one fatigues. Intense efforts rely predomi- first. Increasing frequency of neuronal action potentials
nantly on fast-twitch fibers. These are generally larger excites the muscle again before it has time to relax
than the slow-twitch fibers and belong to larger motor because the twitch is usually much longer than the action
units, so that these are recruited last. This makes subjec- potential. Thus muscle force can summate when excita-
tive sense, because these large muscle fibers in large tion frequency exceeds 1/t, where t is the period of the
motor units increase force in the greatest increments twitch. The tetanic frequency is the frequency at which
when they are recruited. These fast-twitch fibers are also all waviness disappears from the force record. Typically
more easily fatigued than the slow-twitch fibers. Fatigue tetanic force is about five times the twitch force.
of slow-twitch muscles takes longer to produce. Recruitment is usually the most important way of grad-
Therefore, it makes sense that these slow-twitch fibers, ing muscle force and can be responsible for a 100-fold or
which are smaller and belong to smaller motor units, greater range of muscle force.
are recruited early on and so are active almost every Stretching a muscle produces a passive force that
time the muscle is activated, even for tasks requiring lit- increases nonlinearly with stretch. The active force,
tle force. The differences in fatiguability of different the increment of force caused by excitation, changes
muscles led Burke to propose a system of classification biphasically with muscle length. It increases with increas-
of muscles based on four types: ing muscle length when the muscle is short, reaches
1. S 5 slow-twitch fibers a maximum, and then decreases with further stretch of
2. FR 5 fast, fatigue resistant the muscle.
3. FI 5 fast, intermediate fatigue resistant The velocity of muscle shortening depends on the
4. FF 5 fast, fatiguable. load. Maximal velocity occurs at zero load, and maxi-
Most muscles consist of thousands of muscle fibers. mum force develops at zero velocity. The power
Most muscles contain all of the different fiber types, but output of muscle varies biphasically with muscle force,
they differ in their relative number. The soleus muscle being maximal at about 1/3Fmax. Both fast- and slow-
in the human consists predominantly of slow-twitch twitch muscles show this behavior. Power output also
muscle fibers, whereas the gastrocnemius consists varies with velocity, being maximal at about 1/3Vmax.
mainly of fast-twitch fibers. However, there is consider- During rapid movements, almost all power derive
able individual variation in the fiber types that are pres- from fast-twitch muscle.
ent in individual muscles. Transformation of muscle Muscle activation can produce force while the muscle
types appears to be limited and obeys a strict progres- shortens, and this is called a concentric contraction.
sion. Increased use of a muscle invariably tends to con- Isometric contractions refer to the activation of muscle
vert muscle fibers into more fatigue-resistant fibers, but under conditions in which it does not change length.
conversion of fast fatigable fibers all the way to slow- In some cases, muscle activation develops a force
twitch fibers does not occur. while the muscle lengthens. This is an eccentric
contraction. Concentric contractions are used to accel-
erate objects or body parts. Isometric contractions are
SUMMARY used to fix joints in some configuration, whereas
Skeletal muscles consist of thousands of muscle fibers eccentric contractions are used to decelerate objects
which are large multinucleated cells. Each muscle or body parts.
fiber is controlled by a single motor neuron that Muscle fibers often are oriented at an angle with respect
forms a neuromuscular junction approximately in the to the direction of muscle action. This is called the pin-
middle of the fiber. Motor neurons branch and form nation angle. This allows more muscle fibers to fit into
junctions with one-to-many muscle fibers. All of the the volume occupied by the muscle and increases force
muscle fibers innervated by a single motor neuron but decreases muscle velocity.
constitute the motor unit. Large motor units typically
have large motor neurons in the spinal cord, and Fatigue is the loss of muscle force due to prolonged use.
small motor units have small motor neurons. The There are two types of fatigue: rapid onset of fatigue
small motor neurons are more excitable and are brought about by continuous maximal stimulation and
recruited first. slower onset of fatigue brought about by repetitive but
submaximal activation of the muscles.
A single action potential on a motor neuron produces a
twitch. Brief twitches characterize fast-twitch muscles: Muscles can be classified on the basis of their contractile
they develop force rapidly and relax rapidly. Slow-twitch activities. Thus there are slow-twitch fibers, fast and
fibers develop force more slowly and relax more slowly. fatigue-resistant fibers, fast intermediate (with respect to
Muscle force can be graded three ways: (1) by recruiting fatigue) fibers, and fast fatiguable fibers.
304 QUANTITATIVE HUMAN PHYSIOLOGY: AN INTRODUCTION

REVIEW QUESTIONS 5. What is active tension? What is passive tension?

How do they vary with muscle length? Is this an
1. What is a muscle twitch? What is tetany? important way of varying muscle strength?
2. Name three ways that the body varies muscle 6. What is the relationship between force and veloc-
force. Are all equally important physiologically? ity? Is a negative velocity of muscle contraction pos-
3. What is a motor unit? What is recruitment? sible? If so, what do you call such a contraction?
How is recruitment usually affected? How 7. What do we mean by “fast-twitch” and “slow-
much stimulation can be achieved through twitch” fibers? Why is muscle power higher in
recruitment? fast-twitch than in slow-twitch fibers?
4. What is tetany? What is the most rapid train of 8. What are concentric, isometric, and eccentric con-
twitches possible without increasing force? By tractions? What are these used for?
how much can force be increased by increasing 9. What advantage does fiber pinnation afford?
the frequency of stimulation? What is its disadvantage?