CHAPTER

Muscle structure and

function
11
­INTRODUCTION
Learning about the structure and function of muscle, especially skeletal muscle,
is essential for understanding the practical significance of its food product, meat.
Furthermore, muscles are a highly sophisticated system in the living animal designed
for movement (skeletal), digestion (smooth), and respiration (heart). It is also impor-
tant to recognize the role of skeletal muscle after its conversion to meat and subse-
quent entrance into domestic and global markets.

­SKELETAL, SMOOTH, AND CARDIAC MUSCLE

Three classes of muscle can be distinguished functionally and anatomically in the
living animal: skeletal, smooth, and cardiac muscles. Skeletal muscle is defined as
muscles attached to the skeleton. It is the most abundant of the three kinds, making up
about 40% of the bodyweight of animals. Skeletal muscle is responsible for the move-
ment system of the body and functions under voluntary control by the nervous system.
Skeletal muscle cells are multinucleated (100–200 nuclei per cell). Smooth muscle
cells are mononucleated. Smooth muscle is found primarily in the walls of the digestive
tract, viscera, reproductive tract, and blood vessels and is under involuntary nervous
control. Cardiac muscle is found only in the heart and is controlled by the involuntary
nervous system. When observed microscopically, another distinguishing characteristic
of skeletal and heart muscle is the regular array of striations, whereas smooth muscle
is nonstriated. Cardiac muscle cells usually contain only one nucleus and are separated
by intercalated disks that aid in the synchronized contraction of regions of the heart.

­SKELETAL MUSCLE CHARACTERISTICS

Muscles are classified as organs and as such have functional components like other
organs. Fig. 11.1 depicts the organizational structure of a skeletal muscle.
The diagram shows a cross section of muscle enveloped by the connective tis-
sue layer called the epimysium. The figure also includes the muscle organization of
(1) muscle fascicles (bundles) surrounded by the connective tissue layer called the
perimysium and (2) the muscle fibers (individual muscle cells) surrounded by the
connective tissue layer called the endomysium, shown in greater detail in Fig. 11.2.
The Science of Animal Growth and Meat Technology. https://doi.org/10.1016/B978-0-12-815277-5.00011-1 175
© 2019 Elsevier Inc. All rights reserved.
176 CHAPTER 11 Muscle structure and function

FIG. 11.1
A schematic representation of a skeletal muscle in cross section showing the organizational
structure of the muscle bundles (fasciculi) and fibers with their respective surrounding
layers of connective tissue.
Diagram by Marley Dobyns, Animal Science Department, Iowa State University.

FIG. 11.2
A photomicrograph of a cross section of skeletal muscle showing muscle fibers (circled)
within a muscle bundle surrounded by perimysial connective tissue.
Picture courtesy of the Muscle Biology Group, Department of Animal Science, Iowa State University.
 ­Skeletal muscle characteristics 177

The connective tissue layers are thicker in muscles used for power compared with
muscles used for coordination. Also, these thicker layers (sheaths) of connective tis-
sue found in power muscles contribute to less tender meat.
Branching off and continuous with the epimysium is a layer of connective tissue
surrounding the muscle bundles (fasciculi) called the perimysium. An example is
shown in Fig. 11.1. There are both primary and secondary bundles, and the larger
the fasciculi, the coarser the texture of the muscle. Power muscles performing large
movements (legs) have larger bundles (coarser texture) relative to smaller muscles
performing fine movement (back). The perimysium also contains and envelops blood
vessels and nerves. The structured location of another layer of connective tissue,
termed the endomysium, is shown in Fig. 11.1 and branches from the perimysium.
This is the thin layer of connective tissue that loosely surrounds muscle fibers (cells),
the physiological unit of skeletal muscles. Therefore from the myotendinal junction,
collagenous fibers from the tendon connect directly to the muscle cell membrane.
The endomysium is adjacent to the sarcolemma, the cell membrane. The sarcolemma
is the muscle cell membrane responsible for the transfer of chemicals and conduction
of the electrical impulse necessary for contraction. Also, you can observe in the cross
section of muscle other constituents such as blood vessels, capillaries, and inter- and
intramuscular fat.

­MUSCLE FIBERS
Muscle fibers (cells) are the basic unit of muscle. They are long, cylindrical, tubu-
lar cells with tapering conical ends, unbranching, and not perfectly round in cross
section. The number of fibers varies from 50 to 300 in each fasciculus (bundle).
Power muscles have larger fasciculi and larger fibers but fewer fibers per bundle.
The length of muscle cells can range from 1 mm to 34 cm (a centimeter is 0.4 of an
inch and a millimeter is 0.1 cm), although long cells are rare; most of them range
from 1 to 40 mm with an average of 20 to 30 mm. The diameter of muscle cells can
vary from 10 to 100 μm (μm), or microns, because cells taper at each end. Fiber
diameter is affected by many factors. For example, fiber diameters are larger in fish
than in mammals, and fibers in mammals are larger than those in birds. Shorter,
thicker muscles have fibers that are larger in diameter than longer, thinner muscles.
Exercise, increased maturity, and higher plane of nutrition all contribute to the fi-
ber profile. Increased aerobic activity will increase capillary density, mitochondria
number, and even myoglobin content. Increased anaerobic activity (sprinting, lift-
ing weight) will increase fiber diameter and increase the proportion of fast-twitch
fibers. Of course, genetics can influence this development. It is easy to see that
cattle breeds that were once used for draft have a larger frame and often a differing
profile of fibers. This is because of numerous generations of selection for strength
and endurance.
Skeletal muscle is made up of different fiber types dependent on their visual
appearance, metabolism, speed, or strength of contraction. From a gross morpho-
logical classification, fibers can be characterized by their color (red or white) in the
muscle (see Fig. 9.6). Red muscle fibers, found in the endurance muscles of the leg
178 CHAPTER 11 Muscle structure and function

and shoulder, appear red because they have great concentrations of myoglobin (the
pigment in muscle and meat). They are further characterized by their long, slow,
sustained contractile activity and by their high oxidative (aerobic) enzyme activity.
Thus red muscle fibers have a greater concentration of mitochondria associated with
their oxidative activity. Conversely, those fibers with low concentrations of myoglo-
bin found in muscles such as porcine longissimus (loin) or poultry pectoralis major
(breast) muscles are termed white muscle fibers. These fibers contract quickly and
for a short duration. For this type of contractile activity, white fibers are dependent
upon glycolytic metabolism (anaerobic), large stores of glycogen (noted sometimes
as glycogen granules), rapid rates of glycolysis, and a highly developed sarcoplas-
mic reticulum as depicted in Fig. 11.3. The sarcoplasmic reticulum is a network of
membranous tubules that have the ability to bind and release Ca2+ and are involved in
the mechanism of muscle contraction. Intermediate fibers have properties in between
those of red and white fibers, hence the name intermediate. Indeed, few muscles
contain muscle cells of just one fiber type.
Based on speed of contraction as a classification, slower contracting fibers are
classified as Type I, whereas faster contracting fibers are characterized as Type II.

FIG. 11.3
A diagrammatic representation of the sarcoplasmic reticulum within a skeletal muscle
fiber. The cell membrane has been peeled back to show the sarcoplasmic reticulum.
Diagram by Marley Dobyns, Animal Science Department, Iowa State University.
 ­Skeletal muscle characteristics 179

Other classifications of fiber types are more complex and sophisticated. That is,
classification can be based on their inherent glycolytic or oxidative metabolism,
and myosin ATPase activity. With the use of histochemical analysis, fiber types re-
act differently to certain oxidative, glycolytic, and ATPase enzyme-specific stains.
By using the ATPase stain, three different fiber types can be observed: red, white,
and intermediate as described in Table 4.1. Differences in these fiber types can then
be observed, measured, and photographed by using an appropriately equipped light
microscope.

­MYOFIBRILS
Fig. 11.4 provides a schematic of the hierarchal relationship of whole muscle
to a relaxed myofibril. Muscles are organs consisting of muscle fibers (cells)
described in Figs. 11.1 and 11.2. Fibers or cells are specialized in structure and
function in that their structure is highly organized and their function involves
contractile activity.
When observed in the electron microscope, there is a regular array of alternating
dark and light banding patterns (Fig. 11.5). Fibers are made up of myofibrils, and
myofibrils (Fig. 11.4) are subcellular structures specialized in contractile activity.
Myofibrils make up more than 50% of total protein in the cell and are not encased in a
membrane. The proteins constituting the myofibrils are insoluble at the ionic strength
of the cell, and this is the reason that myofibrils can be observed microscopically as
structural entities. Adjacent myofibrils lie with their light and dark bands in register,
and as a result confer a cross-striation appearance upon the entire cell. As observed
in Fig. 11.6, a relaxed myofibril has repetitive bands. Under postmortem conditions,
the relaxed myofibril stage results in more tender meat than the constricted stage.
The dark band, or A band, is anisotropic (birefringent) and hence is dark in the phase
(light) microscope but bright (birefringent) in the polarizing microscope. The light
band, or I band, is isotropic and is light in the phase but dark (nonbirefringent) in the
polarizing microscope.
The dark, thin, vertical line bisecting the I band is called the Z disk (line). The
sarcomere is the structure from one Z line to the next within the myofibril (Fig. 11.4).
And this distance in a relaxed sarcomere is 2.4–2.8 μm in muscle from almost all do-
mestic species. When muscle contracts, sarcomere lengths decrease to 2 μm, or even
less. The myofibril is a series of several hundred to thousands of sarcomeres within
the muscle cell, depending on the length of the muscle cell.
A light area in the middle of the A band is called the H (heller) zone, observed
only when the myofibril is in a relaxed state. Myofibrils are 1–3 μm in diameter and
may occasionally branch. The M line in the center of the H zone, the pseudo H zone,
and thick, thin, titin, and nebulin filaments can only be observed for details with the
electron microscope. Table 11.1 provides information about the components mak-
ing up the microscopic appearance of the myofibril. The abbreviations using letters
for these components come from German words used by early microscopists, for
example, Z line (disk) = zwischenscheibe.
180 CHAPTER 11 Muscle structure and function

Skeletal muscle organization

Muscle

Muscle bundle

Muscle fiber
single fiber
muscle cell Myofibril
myofiber

Sarcomere

A-band Z-line M-line I-band

Thin filament
Nebulin
Troponin Nonoverlap
Actin
region

Thin filament
Tropomyosin
Overlap region Titin
Thick filament
Nebulin
Myosin

C-protein (MBP-C)

Tropomodulin
FIG. 11.4
A schematic representation of the organization of components of a skeletal muscle down to
the contractile elements of a myofibril.
Diagram courtesy of the American Meat Science Association, drawing by Darl Ray.

­SUMMARY
The purpose of this chapter is to show the importance and relationship of the struc-
ture and function of living and postmortem muscle. The knowledge, understanding,
and application of muscle structure and function to postmortem muscle are funda-
mental in securing the quantitative and qualitative characteristics desired in meat and
meat products. The structure of muscle is presented to show the gross anatomical
 ­Summary 181

FIG. 11.5
An electron micrograph of a longitudinal section of a skeletal muscle myofibril showing
the contractile proteins and distinctive Z lines. Note the intricate assembly and the
arrangement of adjacent myofibrils that allows for coordinated contraction within fibers
and, eventually, muscle bundles and muscles.
Photograph courtesy of the Muscle Biology Group, Department of Animal Science, Iowa State University.

FIG. 11.6
An electron photomicrograph of a longitudinal section of a myofibril from skeletal muscle
with specific areas and structures labeled.
Photograph courtesy of the Muscle Biology Group, Department of Animal Science, Iowa State University.

Table 11.1 Definition of terms used to describe the microscopic properties

and appearance of striated skeletal muscle
Term Definition

A band Anisotropic, birefringent (bright) in polarizing, dark in phase

I band Isotropic, nonbirefringent (dark) in polarizing, bright (light) in phase
Z line (disk) Zwischenscheibe (between disks)
M line Mittelscheibe (middle disk)
H zone Heller (bright)
182 CHAPTER 11 Muscle structure and function

parts as well as the microscopic elements of muscle and their relationship to muscle
contraction-relaxation and to postmortem meat quality characteristics, especially
tenderness. The function of living muscle is that of contractile activity in the move-
ment system. The remarkable discovery of thick and thin filaments and their interac-
tion resulting in contraction-relaxation has proven to be valuable in meat science and
the meat industry.
Postmortem skeletal muscle serves as a food, meat. Many paths and processes
are taken to transform the carcass of an animal into wholesale and retail fresh and
processed meat items for eventual consumer purchase. These items can take on the
form of tender and less tender and cured and smoked retail cuts, ground meat, hot
dogs, and jerky. The large number of different meat products is designed to address
income level, ethnic background, and regional preferences and offers convenience,
variety, texture, and taste.

­QUESTIONS FOR STUDY AND DISCUSSION TOPICS

1. Name the three layers of connective tissue found in skeletal muscle.
2. Compare the structural similarities and differences of the three types of muscle.
3. What is the contractile unit of skeletal muscle and how long is this unit in
relaxed muscle?
4. Describe the types of muscle fibers found in skeletal muscle and provide
examples where each may be found.
5. Draw the structure of a myofibril in longitudinal section and label the unique
areas as seen in the electron microscope.
6. Where are the mitochondria located within the muscle fiber?