10/24/2023

Excitable Tissue: Muscle

Lecture 4
2nd Year Pharmacy Students
Munaf S. Saleh
Ph.D. Pharmacology

Introduction
• Muscle cells, like neurons, can be excited chemically, electrically, and
mechanically to produce an AP that is transmitted along their cell
membranes.
• Unlike neurons, they respond to stimuli by activating a contractile
mechanism. The contractile protein myosin and the cytoskeletal
protein actin are abundant in muscle, where they are the primary
structural components that bring about contraction.
• Muscle is divided into three types: skeletal, cardiac, and smooth
• About 40% of the body is skeletal muscle, and perhaps another 10% is
smooth and cardiac muscle

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Introduction
• The three muscle types use common principles to generate force
• Contraction is initiated by a rise in intracellular free Ca+2 conc which
facilitates interaction between actin and myosin that slide against each other
using ATP to generate force
• Sliding cause muscle cells and the myofilaments to contract and shorten
• Although the mechanisms by which force is generated is similar in the
three muscle types, there are significant differences in their
organization and in the way they initiate and control the contractile
force

SKELETAL MUSCLE (SM) MORPHOLOGY

• ORGANIZATION
• SM consists of individual muscle fibers (MFs) that
are the “building blocks” of the muscular system
• Each MF is a single cell that is multinucleated, long,
cylindrical, and surrounded by a cell membrane,
the sarcolemma.
• MFs consist of myofibrils, which are divisible into
individual filaments. These myofilaments contain
several proteins that together make up the
contractile machinery of the SM.
• SM contractile mechanism depends on the
proteins myosin-II, actin, tropomyosin, and
troponin
• Troponin is made up of three subunits:
troponin I, troponin T, and troponin C.

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STRIATIONS
• The parts of the cross-striations are frequently identified by letters.
• The light I band is divided by the dark Z line, and the dark A band has
the lighter H band in its center.
• A transverse M line is seen in the middle of the H band.
• The area between two adjacent Z lines is called a sarcomere

STRIATIONS
• The thick filaments (twice the
diameter of the thin filaments) are
made up of myosin
• The thin filaments are made up of
actin, tropomyosin, and troponin.
• Thick filaments are lined up to
form the A bands, whereas the
array of thin filaments extends out
of the A band and into the I bands
• The Z lines allow for anchoring of
the thin filaments

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STRIATIONS
• The form of myosin found in muscle is myosin-II,
with two globular heads.
• One of these heads form cross-bridges with actin at any
given time. These heads contain an actin-binding site and
a catalytic site that hydrolyzes ATP.
• The thin filaments are polymers made up of two
chains of actin that form a long double helix.
• Tropomyosin molecules are long filaments located in
the groove between the two chains in the actin
• Each thin filament contains 300–400 actin molecules
and 40–60 tropomyosin molecules.

STRIATIONS
• Troponin molecules are small globular units located
at intervals along the tropomyosin molecules
• Each of the three troponin subunits has a unique
function:
• Troponin T binds the troponin components to
tropomyosin;
• troponin I inhibits the interaction of myosin with actin
• troponin C contains the binding sites for the Ca 2+ that
helps to initiate contraction.
• A summary of the structural organization in SM:
Muscle → MF → Myofbril → Thick filaments
(Myosin)→ Thin filaments (Actin, Troponin,
Tropomyosin)

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STRIATIONS
• Additional structural proteins that are important
in SM function include
1. Actinin binds actin to the Z lines.
2. Titin: connects the Z lines to the M lines
• It provides muscle with its elasticity.
• These springy titin molecules act as a framework that
holds the myosin and actin filaments in place.
• One end of the titin molecule is elastic and is attached
to the Z disk, acting as a spring and changing length as
the sarcomere contracts and relaxes
3. Desmin adds structure to the Z lines in part by
binding the Z lines to the plasma membrane.

SARCOTUBULAR SYSTEM
• The muscle fibrils are surrounded by vesicles and tubules
made up of membranes
• These structures form the sarcotubular system, which is
made up of a T system and a sarcoplasmic reticulum (SR).
• The T system of transverse tubules, which is continuous
with the sarcolemma of MF, forms a grid perforated by the
individual muscle fibrils.
• The T system provides a path for the rapid transmission of AP from
the cell membrane to all muscle fibrils
• The SR, which forms an irregular curtain around each of
the fibrils, has enlarged terminal cisterns in close contact
with the T system at the junctions between the A and I
bands.
• The SR is an important store of Ca 2+ and also participates in
muscle metabolism.

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ELECTRICAL CHARACTERISTICS OF SM
• The electrical events in SM and the ionic fluxes that underlie them
share distinct similarities to those in nerve, with quantitative
differences in timing and magnitude.
• The RMP of SM is about –90 mV. The AP lasts 2–4 ms and is
conducted along the MF at about 5 m/s.
• The absolute refractory period is 1–3 ms long, and the after-
polarizations, with their related changes in threshold to electrical
stimulation, are relatively prolonged.
• As in nerves, depolarization is largely a manifestation of Na + influx,
and repolarization is largely a manifestation of K + efflux.

CONTRACTILE RESPONSES
• It is important to distinguish between the electrical and mechanical
events in skeletal muscle.
• Although one response does not normally occur without the other,
their physiologic bases and characteristics are different.
• MF membrane depolarization normally starts at the motor end plate,
the specialized structure under the motor nerve ending.
• The AP is transmitted along the MF and initiates the contractile
response.

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Neuromuscular Junction and

the Motor End plate
• MFs are innervated by large myelinated nerve
fibers that originate from motoneurons in the
the spinal cord.
• Each nerve ending makes a junction, called
the neuromuscular junction, with the MF
near its midpoint
• The nerve fiber forms a complex of branching
nerve terminals that invaginate into the
surface of the MF but lie outside the MF
plasma membrane. The entire structure is
called the motor end plate

THE MUSCLE TWITCH

• A single AP causes a brief contraction followed by
relaxation. This response is called a muscle twitch.
• The twitch starts about 2 ms after the start of
depolarization of the membrane, before repolarization
is complete.
• The duration of the twitch varies with the type of
muscle being tested.
• “Fast” MFs: those concerned with fine, rapid, precise
movement, have twitch durations as short as 7.5 ms.
• “Slow” MFs: those involved in strong, gross, sustained
movements, have twitch durations up to 100 ms.

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GENERAL MECHANISM OF MUSCLE CONTRACTION

1. AP causes the discharge of the motor neuron and the release of the
neurotransmitter acetylcholine (Ach) at the motor end plate
2. Binding of Ach to nicotinic Ach receptor
3. Increased Na+ and K+ conductance in end-plate membrane and generation
of end-plate AP
4. Generation of AP in MFs
5. Inward spread of depolarization along T tubules
6. Release of Ca2+ from terminal cisterns of SR and diffusion to thick and thin
filaments
7. Binding of Ca2+ to troponin C, uncovering myosin-binding sites on actin
8. Formation of cross-linkages between actin and myosin and sliding of thin
on thick filaments, producing movement

MOLECULAR BASIS OF CONTRACTION

• The process by which muscle contraction is brought about is a sliding of the
thin filaments over the thick filaments.
• This sliding occurs when the myosin heads bind firmly to actin, bend at the junction
of the head with the neck, and then detach. This “power stroke” depends on the
simultaneous hydrolysis of ATP
• In resting muscle, troponin I is bound to actin and tropomyosin and covers
the sites where myosin heads interact with actin.
• Following an AP, cytosolic Ca 2+ is increased and free Ca 2+ binds to
troponin C.
• This binding results in a weakening of the troponin I interaction with actin and
exposes the actin binding site for myosin to allow for formation of myosin/actin
cross-bridges

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MOLECULAR BASIS OF CONTRACTION

A. At rest, myosin heads are bound to ADP and are said to be in
a “cocked” position in relation to the thin filament, which
does not have Ca 2+ bound to the troponin–tropomyosin
complex.
B. Ca 2+ bound to the troponin–tropomyosin complex induces a
conformational change in the thin filament allowing myosin
heads to cross-bridge with actin.
C. Myosin heads rotate, move the attached actin and shorten
the MF, forming the power stroke.
D. At the end of the power stroke, ATP binds to a now exposed
site, and causes a detachment from the actin filament.
E. ATP is hydrolyzed into ADP and inorganic phosphate (P i ) and
this chemical energy is used to “re-cock” the myosin head
• As long as Ca 2+ remains elevated and sufficient ATP is
available, this cycle repeats.
• Many myosin heads cycle at or near the same time, and they
cycle repeatedly, producing gross muscle contraction.

Excitation–contraction coupling
• It is the process by which depolarization of MF
initiates contraction
• The AP is transmitted to all the fibrils in the fiber
via the T system. It triggers the release of Ca 2+
from the SR next to the T system (HOW?).
• Depolarization of the T tubule membrane
activates the SR via dihydropyridine receptors
(DHPR)
• DHPR are voltage-gated Ca channels in the T tubule
2+

membrane
• The DHPR unlocks release of Ca 2+ from the
nearby SR via physical interaction with the
ryanodine receptor (RyR)

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Excitation–contraction coupling.
• Ca 2+ is reduced in the muscle cell by the sarcoplasmic or
endoplasmic reticulum Ca 2+ ATPase (SERCA).
• This pump uses ATP to remove Ca 2+ from the cytosol back into the SR, where
it is stored until released by the next AP.
• Once the Ca 2+ concentration outside the SR has been lowered
sufficiently, chemical interaction between myosin and actin ceases
and the muscle relaxes.
• Note that ATP provides the energy for both contraction (at the myosin
head) and relaxation (via SERCA).

Steps in relaxation
• Ca2+ pumped back into sarcoplasmic reticulum
• Release of Ca2+ from troponin
• Cessation of interaction between actin and myosin

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Sources of ATP for Muscle Contraction

• The first source that is used to reconstitute
the ATP is phosphorylcreatine which is
instantly cleaved, and its released energy
causes bonding of a new phosphate ion to
ADP to reconstitute the ATP.
• However, the total amount of
phosphorylcreatine in the MF is small
• The combined energy of both the stored
ATP and the phosphorylcreatine in the
muscle is capable of causing maximal
muscle contraction for only 5 to 8 seconds

Sources of ATP for Muscle Contraction

• The second source of energy used to reconstitute both ATP and
phosphorylcreatine, is glycolysis—the breakdown of glycogen
previously stored in the muscle cells.
• Rapid enzymatic breakdown of the glycogen to pyruvic acid and lactic acid
liberates energy that is used to convert ADP to ATP (faster than oxidative
metabolism)
• Occur even in the absence of oxygen, so muscle contraction can be sustained
for many seconds and sometimes up to more than 1 minute
• The accumulation of lactic acid lowers the pH within the muscle. The change
in pH may ultimately alter the activity of enzymes involved with energy
production as well as cross bridge cycling

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Sources of ATP for Muscle Contraction

• The third and final source of energy is oxidative
metabolism (combining O2 with the end products
of glycolysis and with various other cellular
foodstuffs to liberate ATP)
• The foodstuffs are CHO, fats, and protein.
• More than 95% of all energy used by the muscles
for sustained long-term contraction is derived
from oxidative metabolism
• For extremely long-term maximal muscle
activity—over many hours—the greatest energy
proportion comes from fats but, for periods of 2
to 4 hours, as much as one half of the energy can
come from stored CHO

TYPES OF CONTRACTION
• It is possible for muscle contraction to occur without an appreciable
decrease in the length of the whole muscle
• Such a contraction is called isometric (“same measure” or length).
• For example, supporting an object in a fixed position, such as carrying a book
or a backpack, requires isometric contraction.
• This contraction also occurs when attempting to move an object that is too
heavy to shift or reposition.
• In this case, the muscle may exert maximal force against the object; however, because
the object does not move, the length of the contracting muscle does not change

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TYPES OF CONTRACTION
• Contraction against a constant load with a decrease in muscle length
is isotonic (“same tension”).
• For example, when lifting an object, the muscle contracts and becomes
shorter while the weight of the object remains constant.
• Isotonic contractions are performed for movement of the body, such as
moving the legs when walking

TYPES OF CONTRACTION
• Many activities require both types of contractions by the muscles
• An example is running
• When one of the legs hits the ground, isometric contraction of the muscles
within this limb keep it stiff and help to maintain body support
• At the same time, isotonic contractions in the opposite leg move it forward to
take the next stride

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Summation of Contraction
• The MF electrical response to repeated stimulation is like that of nerve
• MF is electrically refractory only during the rising phase and part of the
falling phase of the spike potential.
• At this time, the contraction initiated by the first stimulus is just
beginning.
• However, because the contractile mechanism does not have a refractory period,
repeated stimulation before relaxation has occurred produces additional
activation of the contractile elements and a response that is added to the
contraction already present.
• This phenomenon is known as summation of contractions.
• The tension developed during summation is considerably greater than
that during the single muscle twitch.

Summation of Contraction
• With rapidly repeated stimulation, activation of the contractile
mechanism occurs repeatedly before any relaxation has occurred, and
the individual responses fuse into one continuous contraction. Such a
response is called a tetanus (tetanic contraction).
• It is a complete tetanus when no relaxation occurs between stimuli
and an incomplete tetanus when periods of incomplete relaxation
take place between the summated stimuli.
• During a complete tetanus, the tension developed is about four times
that developed by the individual twitch contractions.

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Cardiac Muscles
(CM)

CARDIAC MUSCLE MORPHOLOGY

• The striations in CM are similar to those in SM, and Z lines are present.
• Large numbers of elongated mitochondria are in close contact with the
muscle fibrils.
• The MFs branch and interdigitate, but each is a complete unit surrounded
by a cell membrane.
• Where the end of one MF abuts on another, the membranes of both
fibers parallel each other through an extensive series of folds.
• These areas, which always occur at Z lines, are called intercalated disks.
• They provide a strong union between fibers, maintaining cell-to-cell cohesion, so
that the pull of one contractile cell can be transmitted along its axis to the next.

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CM MORPHOLOGY
• Along the sides of the MF next to the
disks, the cell membranes of adjacent
fibers fuse for considerable distances,
forming gap junctions.
• These junctions provide low-resistance
bridges for the spread of excitation
from one fiber to another.
• They permit CM to function as if it were a
syncytium
• The T system in CM is located at the Z
lines rather than at the A–I junction,
where it is located SM.

ELECTRICAL PROPERTIES: RMP & AP

• The RMP of CM cells is about –80 mV.
• Stimulation produces a propagated AP that is
responsible for initiating contraction
• Depolarization proceeds rapidly and an
overshoot of the zero potential is present, as
in SM and nerve, but this is followed by a
plateau before the membrane potential
returns to the baseline.
• Depolarization lasts about 2 ms, but the
plateau phase and repolarization last 200 ms
or more.
• Repolarization is therefore not complete until
the contraction is half over.

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RMP & AP
• The initial rapid depolarization and the overshoot (phase 0)
are due to opening of voltage-gated Na + channels similar to
that occurring in nerve and SM.
• The initial rapid repolarization (phase 1) is due to closure of
Na + channels and opening of one type of K + channel.
• The subsequent prolonged plateau (phase 2) is due to a
slower but prolonged opening of voltage-gated Ca 2+
channels (Ca 2+ entry).
• CM contain T- and L-types of Ca channels, but the Ca current is
2+ 2+

mostly due to opening of the slower L-type Ca channels

2+

• Final repolarization (phase 3) to the RMP (phase 4) is due to

closure of Ca 2+ channels and a slow, delayed increase of K
+ efflux through various types of K + channels

MECHANICAL PROPERTIES
CONTRACTILE RESPONSE
• The contractile response of CM begins just after
the start of depolarization and lasts about 1.5
times as long as the AP. The role of Ca 2+ in
excitation–contraction coupling is similar to its
role in SM.
• However, it is the influx of EC Ca 2+ through the DHPR
in the T system that triggers calcium-induced calcium
release through the RyR at the SR.
• Because there is a net influx of Ca 2+ during
activation, there is also a more prominent role for
plasma membrane Ca 2+ ATPases and the Na +
/Ca 2+ exchanger in recovery of IC Ca 2+
concentrations

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MECHANICAL PROPERTIES
CONTRACTILE RESPONSE
• During phases 0 to 2 and about half of phase 3 (until the membrane
potential reaches approximately –50 mV during repolarization), CM
cannot be excited again; that is, it is in its absolute refractory period.
It remains relatively refractory until phase 4.
• Therefore, tetanus of the type seen in SM cannot occur. Of course,
tetanization of cardiac muscle for any length of time would have
lethal consequences, and in this sense, the fact that CM cannot be
tetanized is a safety feature.

ISOFORMS
• CM is generally slow and has relatively low ATPase activity.
• Its fibers are dependent on oxidative metabolism and hence on a
continuous supply of O 2 .
• The human heart contains both the α and the β isoforms of the myosin
heavy chain (αMHC and β MHC).
• β MHC has lower myosin ATPase activity than α MHC.
• Both are present in the atria, with the α isoform predominating, whereas the β
isoform predominates in the ventricle.
• The spatial differences in expression contribute to the well-coordinated
contraction of the heart.

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CORRELATION BETWEEN MUSCLE FIBER LENGTH & TENSION

• The correlation in CM is similar to that in SM; there

is a resting length at which the tension developed
on stimulation is maximal.
• The initial fiber length is determined by the degree
of diastolic filling of the heart, and the pressure
developed in the ventricle is proportional to the
volume of the ventricle at the end of the filling
phase (Starling’s law of the heart).
• The developed tension increases as the diastolic volume
increases until it reaches a maximum, then tends to
decrease. The decrease is due to beginning disruption of
the myocardial fibers

CORRELATION BETWEEN MUSCLE FIBER LENGTH & TENSION

• The CM contraction force can be also increased by catecholamines

without a change in muscle length.
• This positive ionotropic effect is mediated via innervated β 1 –adrenergic
receptors, cyclic AMP (cAMP), and their effects on Ca 2+ homeostasis.
• The heart also contains noninnervated β 2 -adrenergic receptors, which also
act via cAMP, but their ionotropic effect is smaller and is maximal in the atria.
• cAMP activates protein kinase A, and this leads to phosphorylation of the
voltage dependent Ca 2+ channels, causing them to spend more time in the
open state. cAMP also increases the active transport of Ca 2+ to the SR, thus
accelerating relaxation and consequently shortening systole.
• This is important when the cardiac rate is increased because it permits adequate
diastolic filling

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METABOLISM
• Mammalian hearts have an abundant blood supply, numerous mitochondria,
and a high content of myoglobin, a muscle pigment that can function as O2
storage mechanism.
• Normally, less than 1% of the energy is provided by anaerobic metabolism.
• During hypoxia, this figure may increase to nearly 10%; but under totally anaerobic
conditions, the energy liberated is inadequate to sustain ventricular contractions.
• Under basal conditions, 35% of the caloric needs of the heart are provided by
CHO, 5% by ketones and amino acids, and 60% by fat.
• However, the proportions of substrates utilized vary greatly with the
nutritional state.
• After ingestion of large glucose amounts, more lactate and pyruvate are used; during
prolonged starvation, more fat is used. Circulating free FA normally account for almost
50% of the lipid utilized.

Smooth Muscles (SMS)

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SMOOTH MUSCLE MORPHOLOGY

• SMS is distinguished anatomically from SM and CM because it lacks visible
cross-striations.
• Actin and myosin-II are present, and they slide on each other to produce
contraction
• However, they are not arranged in regular arrays and so the striations are absent.
• Instead of Z lines, there are dense bodies in the cytoplasm and attached to
the cell membrane, and these are bound by α-actinin to actin filaments.
• SMS also contains tropomyosin, but troponin appears to be absent
(calmodulin instead of troponin).
• A SR is present, but it is less extensive than those observed in SM or CM.
• In general, SMS contain few mitochondria and depend, to a large extent, on
glycolysis for their metabolic needs.

TYPES
1. Unitary (or visceral ) SMS
• occurs in large sheets, has many low-resistance
gap junctional connections between individual
muscle cells, and functions in a syncytial fashion.
• Found primarily in the walls of hollow viscera.
• The musculature of the intestine, the uterus, and the
ureters are examples.
• en passant endings of nerve fibers on fewer
cells, with excitation spreading to other cells by
gap junctions

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TYPES
2. Multiunit SMS
• Made up of individual units with few (or no) gap
junctional bridges.
• Found in structures such as the iris of the eye, in
which fine, graded contractions occur.
• It is not under voluntary control, but it has many
functional similarities to SM.
• Each cell has en passant endings of nerve fibers
• In addition, SMS cells respond to hormones
and other circulating substances.
• Blood vessels have both unitary and multiunit
smooth muscle in their walls.

ELECTRICAL & MECHANICAL ACTIVITY

• Unitary SMS is characterized by the instability of its membrane potential
and by the fact that it shows continuous, irregular contractions that are
independent of its nerve supply.
• This maintained state of partial contraction is called tonus, or tone.
• The membrane potential has no true “resting” value, being relatively
low when the tissue is active and higher when it is inhibited, but in
periods of relative quiescence values for resting potential are on the
order of –20 to –65 mV

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MOLECULAR BASIS OF CONTRACTION

• As in other muscles, Ca 2+ plays a prominent role in initiating SMS
contraction. However, the source of Ca 2+ increase can be quite different in
unitary SMS.
• Depending on the activating stimulus, Ca 2+ increase can be due to influx through
voltage- or ligand-gated plasma membrane channels, efflux from intracellular stores
through the RyR or through the inositol trisphosphate receptor (IP 3 R) Ca 2+ channel,
or via a combination of these channels.
• In addition, the lack of troponin in SMS prevents Ca 2+ activation via troponin
binding.
• Rather, myosin in SMS must be phosphorylated for activation of the myosin
ATPase
• Ca 2+ binds to calmodulin, and the resulting complex activates calmodulin-dependent
myosin light chain (MLC) kinase. This enzyme catalyzes the phosphorylation of the MLC
increasing its ATPase activity

MOLECULAR BASIS OF CONTRACTION

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Types of smooth muscle contraction

1.Rhythmic contraction (rhythmicity): a pattern of repeated
contractions, this caused by self- exciting fibers that deliver
spontaneous impulses transmitted from cell to cell
• Rhythmicity is responsible for the wavelike motion called
peristalsis (alternate contraction and relaxation) e.g. in
GIT.

2.Tonic contraction: continuous contraction of smooth

muscle,
• This type is found in the sphincters and wall of blood
vessels.

Relaxation
• Myosin is dephosphorylated by MLC
phosphatase in the cell.
• Relaxation of the muscle presumably
occurs when the Ca 2+ -calmodulin
complex finally dissociates or when
some other mechanism comes into
play

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RELAXATION
• Endothelial cells that line the inside of blood vessels could release a
substance that relaxed smooth muscle (nitric oxide (NO)).
• NO produced in endothelial cells is free to diffuse into the SMS for its
effects.
• Once in muscle, NO directly activates a soluble guanylate cyclase to
produce another second messenger molecule, cyclic guanosine
monophosphate (cGMP).
• This molecule can activate cGMP-specific protein kinases that can affect ion
channels, Ca 2+ homeostasis, or phosphatases, or all of those mentioned,
leading to SMS relaxation

Effects of various agents on the membrane

potential of intestinal smooth muscle

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FUNCTION OF THE NERVE SUPPLY TO SMOOTH

MUSCLE
• The effects of Ach and norepinephrine on unitary SMS serve to
emphasize two of its important properties:
1. its spontaneous activity in the absence of nervous stimulation
2. its sensitivity to chemical agents released from nerves locally or brought to
it in the circulation.
• In mammals, unitary SMS usually has a dual nerve supply from the
two divisions of the autonomic nervous system.
• The function of the nerve supply is not to initiate activity in the muscle but
rather to modify it.
• Stimulation of one division of the autonomic nervous system usually increases
SMS activity, whereas stimulation of the other decreases it.
• In some organs, noradrenergic stimulation increases and cholinergic
stimulation decreases SMS activity; in others, the reverse is true

FORCE GENERATION & PLASTICITY OF SMS

• SMS displays a unique economy when compared to SM.
• Despite approximately 20% of the myosin content and a 100-fold
difference in ATP use when compared with SM, they can generate
similar force per cross-sectional area.
• One of the tradeoffs of obtaining force under these conditions is the
noticeably slower contractions when compared to SM.
• There are several known reasons for these noticeable changes, including
unique isoforms of myosin in SMS and their distinct regulation. The unique
architecture of the SMS also likely contribute to these changes.

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FORCE GENERATION & PLASTICITY OF SMS

• Another special characteristic of SMS is the variability of the tension it
exerts at any given length.
• If a unitary SMS is stretched, it first exerts increased tension. However, if the muscle
is held at the greater length after stretching, the tension gradually decreases.
• Sometimes the tension falls to or below the level exerted before the muscle was
stretched. It is consequently impossible to correlate length and developed tension
accurately, and no resting length can be assigned.
• In some ways, therefore, SMS behaves more like a viscous mass than a
rigidly structured tissue, and it is this property that is referred to as the
plasticity.
• It is the ability of SMS to adapt easily and quickly to length changes without
compromising its ability to generate force

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