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PHYSIOLOGY OF MUSCLE

Muscle cells are “excitable cells”. Like neurons, action potentials can
be produced in response to chemical, electrical, or mechanical stimuli. Due
to the presence of contractile elements, muscle response is able to generate a
force, which could be used in the performance of many activities, for
example, movement of the body parts, mixing of foods, elimination of
wastes, etc.

A. Types of muscles:

There are three types of muscle which differ anatomically and
physiologically from one other.

1. Skeletal muscle
- cross striations are a very prominent feature
- muscle fibers are distinctly separate
- normal contractions is in response to nervous
stimulation
- under voluntary control

2. Cardiac muscle
- is also striated
- functionally responding as a single muscle fiber
- contracts automatically and rhythmically even in the
absence of its nerve supply
- involuntary

3. Smooth muscle
- unstriated
- may posses some automaticity and rhythmicity
- involuntary

B. Muscle contraction:

Muscle contraction is a mechanical change, characterized by
shortening or development of tension and which is associated
with electrical, chemical and thermal changes, all of which are
reversible.

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1. Electrical changes: Excitation of the skeletal muscle fiber.

a) Electrical properties at rest are similar to nerve fibers:
- concentration of K+ higher inside the cell
- cell wall, at rest, is highly permeable to K+ but only
slightly permeable to Na+
- concentration of Na+ higher outside cell
- resting membrane potential is about 90 mv, inside
negative

b) Excitation of the muscle, the action potential.
- the nerve impulse causes release of acethylcholine at
the endings
- at the end plate, an end plate potential is created. If its
magnitude reaches the firing level, an action potential is
produced which moves along the muscle membrane in
both directions. The action potential is transmitted
rapidly to innermost muscle fibers thru the transverse
tubular system. The whole process is termed neuro-
muscular transmission

c) Electrical properties during activity
- the action potential has an overshoot of up to +30 mv.
- duration of an action potential 2-4 millisecond
- rate of conduction through muscle 5 m/sec
- the electrical response always occurs before and is
completed by the time the mechanical response begin

2. Mechanical changes
Depolarization of the muscle fibers results in contraction.
The transformation of electrical to mechanical activity involves
chemical reactions and the process is called excitation-
contraction coupling.

a) The muscle twitch
The response to a single action potential
starts about 2 msec. after the beginning of
depolarization of muscle membrane

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Stages:
(i) Latent period
The interval between the depolarization of the
muscle fiber and the beginning of muscle
contraction.

(ii) Period of Contraction
The muscle fiber contracts and shortens or develop
tensions. The interval between the beginning of the
electrical response to the peak of the tension curve
is sometime called the contraction time.

(iii) Period of relaxation
There is reduction in the developed tension of
force.

. The duration of the muscle twitch varies, depending on the specie and
the type of muscle. It maybe as short as 7.5 msec. in “fast” muscle such as
those concerned with fine, rapid and precise movements. It maybe as long as
100 msec. – 200 msec. in “slow” muscle like those concerned with
maintenance of posture.

b) Types of muscle contraction
(i) Isometric contraction

Contraction without a decrease in the length of the
whole muscle. The muscle exerts tension but does not shorten. No
work is done.

Example:
Contraction of muscles maintaining posture and
equilibrium and when one holds on object.

(ii) Isotonic contraction

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The muscle shortens under a constant load. Weight is
carried through a distance and thus work is performed
(positive work)

If, during the contraction, the tension developed is less
than the opposing force, the muscle is actually stretched
or lengthened (negative work)

( c) Mechanism of shortening
( i) Molecular background

- A myofibril is made up of thick and thin filaments
which contain contractile proteins longitudinally
arranged.
- Each thick filament is surrounded by six thin filaments
which are equally spared in a hexagonal manner
- The thick filament is a bundle of myosin molecules
which are rod-shaped and possesses a double head at
one end. The myosin rods are parallel to each other
while their head are oriented towards both ends and the
filament.
- The thin filament contains the other contractile proteins
– actin, troponin and troypomyosin
- Stands of globular actin molecules form a double helix.
Within the groove of the double helix are found the
tropponin-tropomyosin complex. This complex is
made up of strings of tropomyosin molecules with
troponin molecules located at internvals along it. Each
string of tropomyosin extends over 7 molecules of
actin.
- There are at least there (3) known components of
troponin: Troponin I, interaction inhibitors; Troponin
T, binds the troponin to tropomyosin; Troponin C,
Calcium binding component.

( ii) Excitation- contraction coupling; the process of
contraction

- When the call membrane is depolarized, the action
potential is transmitted to the muscle fibers thru the

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sarcotubular system (T-tubules and sarcoplasmic
reticulum)
- Ca++ ions are released from the terminal cisterns
(lateral saca) of the sarcoplasmic reticulums and
diffuses to the reactive sites.
- Ca++ binds to troponin C
- Removal of inhibitory action of the troponin-
tropomyosin complex on action
- Interaction between actin and myosin; myosin heads
attach to actin reactive sites (formation of cross-
bridges)
- Breakdown of ATP and release of energy
- “Swiveling” or “rowing” movement of myosin heads
attached to actin. After each rowing movement the
myosin head detaches from the actin, swings back and
binds with the next actin reactive site.
- Repeated attachment, swiveling and detachment of the
myosin cause the sliding of the thick and thin filaments
along and past each other thus causing shortening. This
mechanism is the so-caled sliding filament theory of
muscles contraction.

( iii) Relaxation of muscle

- While the cell membrane repolarizes, Ca++ is actively
sequestered back to the cisterns. ATP provides the
energy for this.
- Inhibitory action of troponin-tropomyosin on actin is
re-established.
- Myosin attachments to actin are broken and the
filaments slide back to the pre-contraction position

(d ) The motor unit

An individual motor nerve fiber supplies more than one muscles
Fiber. Each individual motor neuron and all the muscle fiber it
supplies, is called the motor unit. The number of muscles fibers in
the motors unit is variable depending on the function of the
muscle. Muscles which are precisely controlled and rapidly
reacting has smaller number of muscles in the motor unit, like the

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muscles of the hand and eye, where the muscle fibers on the
motor unit maybe as low as 3-6. Muscles concerned with coarser
movements like those for maintainance of posture have more
fibers in the motor unit, averaging 180 fibers.

(3) Chemical Changes:
(a) The immediate source of energy of muscle contraction is ATP.
Hydrolysis of ATP to ADP is associated with the release of a
great amount of energy.
(b) ATP is resynthesized from ADP by the addition of a phosphate
group. Energy from this reaction maybe derived from:

(i) Hydrolysis of phosphoceatine to creatine and phosphates with
liberation of energy
(ii) Breakdown of glucose into pyruvic acid and into CO2 and
water in the presence of oxygen (aerobio glypolysis)
(iii) In the absence of oxygen, breakdown of glucose into pyruvic
acid, then into lactic acid (anerobic glycolysis)
(iv) Oxidation of free fatty acids into CO2 and water.

The energy liberated by the breakdown of carbohydrates is also used
to resynthesize phosphocreatine.

If muscular exertion is to great, the supply of oxygen, even though
markedly increased, becomes insufficient for the aerobic resynthesis
of ATP. Thus the energy provided by the anerobic breakdown of
glucoses to lactic acid is availed of for resynthesis.

The accumulation of lactic acid however reduces pH in the muscles
tissues and inhibits the process of resynthesis.

Thus the oxidative metabolism continues even after the exertion is
over, in order to remove excess, lactic acid as well as to continue the
resynthesis of energy stores. The additional amount of oxygen needed
for these activities is the consequence of the disparity between aerobic
resynthesis and the utilization of energy stores during the increased
muscular effort. Hence, an oxygen debts, is said to have been
incurred.

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4. Thermal changes
The liberation of energy during muscle contraction is associated
with the production of heat.

(a) Resting heat - heat given off by the inactive muscle, it is
due to the metabolic process of the resting muscle.

(b) Initial heat – heat liberated during the active phase of
contraction. It is made-up of the following:

(i) Activation heat - heat production when the muscle
contracts as in a muscle twitch. (in titanic contraction
it is called maintenance heat)
(ii) Hat of shortening - liberated when the muscle
shortens. It is dependent upon the degree of
shortening.

( c ) Recovery heat or delayed heat

Additional amount of hat liberated as a consequence of
metabolic processes which returns the muscles into its original pre-
contraction physical condition.

(i) Heat of relaxation - liberated when isotonic muscle contraction is
over and relaxation occurs. The heat is produced as a result of the
structuring by the load

C. Effects of multiple stimuli on muscle contraction.
1. Treppe (staircase” phenomenon)

If a fresh skeletal muscle is stimulated successively with
maximal stimuli at a frequency which allow the muscle to relax completely,
the tension developed during each twitch increases until the tension becomes
the same during each contraction.

This phenomenon maybe some kind of “warning up” process the
mechanism of which is not well understood.

2. Wave summation and tetanus

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A single maximal stimulus will cause the development maximal
tension. If another stimulus, stronger than maximal, is applied, the twitch
tension developed will still be of the same magnitude.

But if two maximal stimuli are used successively with the second
being applied while the muscle is still responding to the first stimulus,
response greater than that produced by a single maximal stimulus, will be
elicited. Several stimuli applied successively in this manner cause responses
of increasing magnitude. The phenomenon is called wave summation. If
the stimuli will applied at t a rapid rate, all contractile elements are
maximally stimulated, all elastic component are stretched fully and a
sustained contraction is produced. This is called tetanus. If the rate of the
stimulation is slower, mechanical fusion may not occur. The individual
contractions maybe discernible and the phenomenon is called incomplete
tetanus. The rate of stimulation capable the producing tetanus varies.

Examples:
350 stimuli/sec. for internal rectus
30 stimuli/sec. for sole us muscle

The action potential developed in muscle membrane is always of the
same magnitude (all-or-none). The activation of the contractile elements is
not. It is dependent on the amount of Ca++ released from the cisterns.
Successive stimuli produces action potentials which cause more and more
Ca++ to be released, thus increasing the contractile response.

D. Effect of increasing the intensity of the stimulus:
A weak stimulus activates only a few motor units. As the intensity of
the stimulus increase, more and more motor units are activated. At a certain
intensity, more motor units are activated. Any further increase in stimulus
intensity therefore will not increase anymore the magnitude of response.
The process is called recruitment of motor units, also sometime called
quantal summation.

E. Length-Tension Relationship
Muscle is made up of contractile elements and elastic tissues. If no
stretch is initially applied on the muscle, the active tension produced by
muscle contraction will be less since the pull merely stretches the elastic
tissue (series-elastic components). If the muscle will be stretched passively
and passive tension develops, the active tension produced by muscle

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contraction, increases. The greater passive tension, the greater is the tension
produced up to a certain point. Beyond which, any further passive
stretching of the muscle reduce the active tension developed during
contraction. The length of the muscle in which maximum active tension is
developed when the muscle is stimulated is called the resting length. This is
the normal length of the muscle in the body. At thus length, the muscle is
lightly stretched and therefore, there is some degree of passive tension
present. Proof of this, is the shortening of a muscle that occurs when the
tendon is severed from its attachment.

F. Effect of Load on the Velocity of Shortening:
Increasing the load on the muscle slows down the shortening and
reduces the distance of load is moved.

G. Smooth Muscle:
1. Types:
(a) Multi-unit smooth muscle
(i) Characteristics:

- found in ciliary muscles, iris pilomotors and the blood
vessels
- responses somewhat similar to skeletal muscle
- activity controlled by discharge from motor nerves
which is believe to release nor-epinephrine at the
endings.
- Time course of muscle twitch greatly prolonged
- Summation and tetanus maybe produced

(c) Visceral Smooth Muscle
(i) Characteristics
- found in the walls of hollow visceral organs like the
uterus, ureters and intestines.
- Individual fibers are in functional continuity (syncitial)
- Innervated by the fibers of the automatic nervous
system which are responsible for regulation, not the
initiation of muscular activity.
- Involuntary
- Capable of persistent, sustained activity independent of
extrinsic and intrinsic nerve activity. This property is
termed “tonus”.

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- Stretching to a new length initially increases tension but
this tension diminishes as the new length is attained.
This property of “elasticity” allows an increase in the
volume of the organ without a very great rise in
pressure.

(ii) Electrical properties.
- distribution and permeability of K+ across the resting
muscle membrane is similar to other cells,
- resting potential, about 60 mv., is lower and less stable
than that of skeletal muscle.
- Shall fluctuations of the resting potential occurs an if
such a change reaches threshold, an action potential is
produced.

(Additional consideration or smooth muscle physiology will be taken up
ender the different organ systems)

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