muscles
muscles
There are more
than 600 muscles in the human body. Most of
these muscles are attached to the bones of the skeleton bytendons, although a few muscles are attached to
the undersurface of the skin. The primary function of the
muscular system is to move
the skeleton. The
muscle contractions required for movement also
produce heat, which contributes to the maintenance of a
constant body temperature. The other body systems
directly involved in movementare the nervous,
respiratory, and circulatory systems.
The nervous system
transmits the electrochemical impulses that
cause muscle cells to contract. The respiratory system exchanges
oxygen and carbon dioxide between the air
and blood. The circulatory system brings oxygen to
the muscles and takes carbon
dioxide away.These interactions
of body systems are covered in
this chapter,
which focuses on the skeletal muscles.You may recall
from Chapter 4 that there are two other types of
muscle tissue: smooth muscle and cardiac muscle. These
types of muscle tissue will be discussed in other chapters
in relation to the organs of
which they are
part. Before you continue, you may find it helpful to
go back to Chapter 4 and review the structure and
characteristics of skeletal muscle tissue.In this chapter we
will begin with the gross (large) anatomy and
physiology of muscles, then discuss the
microscopic
structure of muscle cells and the biochemistry of muscle
contraction.
MUSCLE STRUCTURE
All muscle cells
are specialized for contraction. Whenthese cells
contract, they shorten and pull a bone tproduce movement.
Each skeletal muscle is made of thousands of
individual muscle cells, which also maybe called muscle
fibers Depending on the work a muscle is required to do, variable
numbers of muscle fibers contract.When picking up a
pencil, for example, only a small
portion of the
muscle fibers in each finger muscle will contract. If the
muscle has more work to do, such as picking up a book,
more muscle fibers will contract to accomplish the
task.Muscles are
anchored firmly to bones by tendons.Most tendons are
rope-like, but some are flat; a flat tendon is called
an aponeurosis.
Tendons are made of fibrous
connective tissue, which, you may
remember, is very
strong and merges with the fascia that covers the
muscle and with the periosteum, the fibrous connective
tissue membrane that covers bones.A muscle usually
has at least two tendons, eachattached to a
different bone. The more immobile or
stationary
attachment of the muscle is its origin; themore movable
attachment is called the insertion. Themuscle itself
crosses the joint of the two bones towhich it is
attached, and when the muscle contracts itpulls on its
insertion and moves the bone in a specific
direction.
MUSCLE
ARRANGEMENTS
Muscles are
arranged around the skeleton so as to bring about a
variety of movements. The two general types of
arrangements are the opposing antagonists and the
cooperative synergists.
Antagonistic
Muscles
Antagonists are
opponents, so we use the term antagonistic muscles for
muscles that have opposing or opposite functions. . The biceps
brachii is the
muscle on the front of the upper arm.The origin of the
biceps is on the scapula (there are actually two
tendons, hence the name biceps), and the insertion is on
the radius. When the biceps contracts,it flexes the
forearm, that is, bends the elbow Recall that when a muscle contracts,
it gets shorter and pulls. Muscles cannot push, for
when they relax they exert no force.Therefore, the
biceps can bend the elbow but cannot straighten it;
another muscle is needed. The triceps brachii is located
on the back of the upper arm. Its origins (the prefix tri
tells you that there are three of
them) are on the
scapula and humerus, and its insertion is on the ulna.
When the triceps contracts and pulls, it extends
the forearm, that is, straightens the elbow.Joints that are
capable of a variety of movements have several sets
of antagonists. Notice how many
ways you can move
your upper arm at the shoulder,
Synergistic
muscles
are those with the same function,or those that work
together to perform a particular function. Recall
that the biceps brachii flexes the forearm. The
brachioradialis, with its origin on the
humerus and
insertion on the radius, also flexes the forearm. There
is even a third flexor of the forearm,the brachialis.
You may wonder why we need three muscles Joints that are
capable of a variety of movements have several sets
of antagonists. Notice how many
ways you can move
your upper arm at the shoulder, for instance.
Abducting (laterally raising) the arm is thefunction of the
deltoid. Adducting the arm is broughtabout by the
pectoralis major and latissimus dorsi. Flexion of the arm
(across the chest) is also a function of the pectoralis
major, and extension of the arm
(behind the back)
is also a function of the latissimus
dorsi. All of
these muscles are described and depicted
in the tables and
figures later in the chapter. Without
antagonistic
muscles, this variety of movements would
not be possible
Muscles may also
be called synergists if they help to
stabilize or
steady a joint to make a more precise
movement possible.
If you drink a glass of water, the
biceps brachii may
be the prime mover to flex the
forearm. At the
same time, the muscles of the shoulder
keep that joint
stable, so that the water gets to
your mouth, not
over your shoulder or down your
chin. The shoulder
muscles are considered synergists
for this movement
because their contribution makes
the movement
effective.
QUESTION: When the
biceps contracts, what happens to its length, and what kind of force
does it exert?
motor areas of the
frontal lobes generate electrochemical
impulses that
travel along motor nerves to muscle fibers,
causing the muscle fibers to contract.For a movement to
be effective, some muscles must contract while
others relax. When walking, for example,antagonistic
muscles on the front and back of the
thigh or the lower
leg will alternate their contractions and relaxations,
and our steps will be smooth and efficient.This is what we
call coordination, and we do not have to think
about making it happen. Coordination takes place below
the level of conscious thought and is regulated by the
cerebellum, which is located below
the occipital
lobes of the cerebrum.
MUSCLE TONE
Except during
certain stages of sleep, most of our muscles are in a
state of slight contraction; this is what is known as muscle
tone. When sitting upright, for example, the tone
of your neck muscles keeps your head up, and the
tone of your back muscles keeps your
back straight.
This is an important function of muscle tone for human
beings, because it helps us to maintain an upright
posture. For a muscle to remain slightly contracted, only a
few of the muscle fibers in that muscle must
contract. Alternate fibers contract so that the muscle as a
whole does not become fatigued. This
is similar to a
pianist continuously rippling her fingers over the keys of
the piano—some notes are always sounding at any
given moment, but the notes that are sounding are
always changing. This contraction of alternate fibers,
muscle tone, is also regulated by the
cerebellum of the
brain.Muscle fibers need
the energy of ATP (adenosine triphosphate) in
order to contract. When they produce
ATP in the process
of cell respiration, muscle fibers also
produce heat. The heat generated by normal muscle tone is
approximately 25% of the total body heat at rest.
During exercise, of course, heat production increases
significantly.
EXERCISE Good muscle tone
improves coordination. When
muscles are
slightly contracted, they can react more rapidly if and
when greater exertion is necessary.Muscles with poor
tone are usually soft and flabby, but exercise will
improve muscle tone.
There are two
general types of exercise: isotonic and isometric. In
isotonic exercise, muscles contract and bring about
movement. Jogging, swimming, and weight lifting are
examples. Isotonic exercise improves muscle tone,
muscle strength, and, if done repetitively against great
resistance (as in weight lifting), muscle
size. This type of
exercise also improves cardiovascular and respiratory
efficiency, because movement exerts demands on
the heart and respiratory muscles.If done for 30
minutes or longer, such exercise may be called aerobic,
because it strengthens the heart and respiratory muscles as well as
the muscles attached to the
skeleton.Isotonic
contractions are of two kinds, concentric
or eccentric. A
concentric contraction is the shorteningof a muscle as it
exerts force. An eccentric contraction is the
lengthening of a muscle as it still exerts force.
Imagine lifting a book straight up (or try it); the triceps
brachii contracts and shortens to straighten the
elbow and raise the book, a concentric contraction. Now
imagine slowly lowering the book.The triceps
brachii is still contracting even as it is lengthening,
exerting force to oppose gravity (which
would make the
book drop quickly). This is an eccentric
contraction.
ENERGY SOURCES FORMUSCLE CONTRACTION
Before discussing
the contraction process itself, let us look first at how
muscle fibers obtain the energy they need to contract.
The direct source of energy for muscle contraction is
ATP. ATP, however, is not stored in large amounts in
muscle fibers and is depleted in a few seconds.The secondary
energy sources are creatine phosphate and glycogen.
Creatine phosphate is, like
ATP, an
energy-transferring molecule. When it is broken down (by an
enzyme) to creatine, phosphate, and energy, the energy
is used to synthesize more ATP. Most of the
creatine formed is used to resynthesize creatine
phosphate, but some is converted to creatinine,a nitrogenous
waste product that is excreted by
the kidneys.The most abundant
energy source in muscle fiber
During strenuous
exercise, the oxygen stored in myoglobin is
quickly used up, and normal circulation may not deliver
oxygen fast enough to permit the completion of cell
respiration. Even though the respiratory rate increases,
the muscle fibers may literally
run out of oxygen.
This state is called oxygen debt, and in this case,
glucose cannot be completely broken down into carbon
dioxide and water. If oxygen is not present (or not
present in sufficient amounts), glucose is converted to an
intermediate molecule called lactic acid, which causes
muscle fatigue.In a state of
fatigue, muscle fibers cannot contract efficiently, and
contraction may become painful. To be in oxygen debt
means that we owe the body some oxygen.Lactic acid from
muscles enters the blood and
circulates to the
liver, where it is converted to pyruvic acid, a simple
carbohydrate (three carbons, about half a glucose
molecule). This conversion requires ATP, and oxygen is
needed to produce the necessary ATP in the liver.
This is why, after strenuous exercise,
the respiratory
rate and heart rate remain high for a time and only
gradually return to normal. Another name proposed for
this state is recovery oxygen uptake, which is a
little longer but also makes sense.Oxygen uptake
means a faster and deeper respiratory
rate. What is this
uptake for? For recovery from strenuous exercise.
MUSCLE FIBER—MICROSCOPIC
STRUCTURE
We will now look
more closely at a muscle fiber, keeping in mind that there
are thousands of these cylindrical cells in one
muscle.Each muscle fiber has its own motor nerve ending;the neuromuscular junction is where the motor
neuron terminates on the muscle fiber.
The axon terminal is the enlarged tip of the motor
neuron; it contains sacs of the neurotransmitter acetylcholine
(ACh). The membrane of the muscle fiber
is the sarcolemma, which contains receptor sites for
acetylcholine, and an inactivator called
cholinesterase. The synapse (or synaptic cleft) is the small space
between the axon terminal and the sarcolemma.Within the muscle
fiber are thousands of individualcontracting units
called sarcomeres, which are arranged end to
end in cylinders called myofibrils. The structure of a
sarcomere is shown in
The Z lines are
the end boundaries of a sarcomere.Filaments of the
protein myosin are in the center of the sarcomere, and
filaments of the protein actin are at the ends,
attached to the Z lines. Myosin filaments are anchored to
the Z lines by the protein titin.Myosin and actin
are the contractile proteins of a muscle fiber.
Their interactions produce muscle contraction.Also present are
two inhibitory proteins, troponin and tropomyosin,
which are part of the actin filaments and
prevent the sliding of actin and myosin
when the muscle
fiber is relaxed.Surrounding the
sarcomeres is the sarcoplasmic reticulum, the
endoplasmic reticulum of muscle cells.The sarcoplasmic
reticulum is a reservoir for calcium ions (Ca[1]2),
which are essential for contraction process.
All of these parts
of a muscle fiber are involved in the contraction
process. Contraction begins when a nerve impulse
arrives at the axon terminal and stimulates the release of
acetylcholine. Acetylcholine generates electrical changes
(the movement of ions) at the sarcolemma of the
muscle fiber. These electrical
changes initiate a
sequence of events within the muscle fiber that is
called the sliding filament mechanism of muscle
contraction. We will begin our discussion with
the sarcolemma.
CONTRACTION—THE
SLIDINGFILAMENT MECHANISM
All of the parts
of a muscle fiber and the electrical changes described
earlier are involved in the contraction process, which is
a precise sequence of events called the sliding
filament mechanism.
In summary, a
nerve impulse causes depolarization of a muscle fiber,
and this electrical change enables the myosin filaments
to pull the actin filaments toward the center of the
sarcomere, making the sarcomere shorter.
All of the sarcomeres shorten and the muscle fiber contracts. A more detailed description of this process is the following:1. A nerve impulse arrives at the axon terminal;
All of the sarcomeres shorten and the muscle fiber contracts. A more detailed description of this process is the following:1. A nerve impulse arrives at the axon terminal;
acetylcholine is
released and diffuses across the
synapse.2. Acetylcholine
makes the sarcolemma more permeable
to Na[1]
ions, which rush into the cell.3. The sarcolemma
depolarizes, becoming negative outside and
positive inside. The T tubules bring the reversal of
charges to the interior of the muscle
cell.4. Depolarization
stimulates the release of Ca[1]2
ions
from the
sarcoplasmic reticulum. Ca[1]2
ions bond to the
troponin–tropomyosin complex, which shifts it away
from the actin filaments.5. Myosin splits
ATP to release its energy; bridges
on the myosin
attach to the actin filaments and pull them toward
the center of the sarcomere, thus making the
sarcomere shorter
6. All of the
sarcomeres in a muscle fiber shorten—the entire muscle
fiber contracts.7. The sarcolemma
repolarizes: K[1]
ions leave the cell, restoring a
positive charge outside and a negative
charge inside. The
pumps then return Na[1]ions outside and K[1]
ions inside.8. Cholinesterase
in the sarcolemma inactivates
acetylcholine.9. Subsequent
nerve impulses will prolong contraction (more acetylcholine
is released).10. When there are
no further impulses, the muscle fiber will relax
and return to its original length.Steps 1 through 8
of this sequence describe a single
muscle fiber
contraction (called a twitch) in response to
a single nerve
impulse. Because all of this takes place
in less than a
second, useful movements would not be possible if muscle
fibers relaxed immediately after contracting.
Normally, however, nerve impulses arrive in a continuous
stream and produce a sustained contraction called tetanus,
which is a normal state not to
be confused with
the disease tetanus. When in tetanus, muscle fibers remain
contracted and are capable of effective
movements. In a
muscle such as the biceps brachii that flexes the
forearm, an effective movement means that many of its
thousands of muscle fibers are in tetanus,a sustained
contraction.
As you might
expect with such a complex process, muscle contraction
may be impaired in many different ways. Perhaps the
most obvious is the loss of nerve impulses to muscle
fibers, which can occur whennerves or the
spinal cord are severed, or when a stroke
References
•Campbell, N.A., J.B.
Reece, L.G. Mitchell, and M.R. Taylor (2003) Biology: Concepts and Connections, Benjamin/Cummings
•Hoar, W.S. (1983) General
and Comparative Physiology, Prentice-Hall
•Snell, R.S.(2003) Clinical Neuroanatomy for
Medical Students.
Lippincott Williams &Wilkins
•Rosenzweig, M.R., S.M. Breedlove, and A.L. Leiman
(2002) Biological
Psychology: An Introduction to Behavioural, Cognitive and Clinical
Neuroscience. Sinauer Associates
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