Muscle Physiology

Human Physiology – From Cells to Systems | 9e
Lauralee Sherwood
8
Muscle Physiology
Prof. Walid Hleihel

Dr. Perla Gerges
2016. All Rights Reserved.Dr. Tania Bitar
© Cengage Learning 2016
The Muscles
o Largest group of tissues in the body
o Constitute approximately half the body’s weight
• Skeletal muscles accounting for
 40% of the body weight in men

 32% of the body weight in women
© Cengage Learning 2016. All Rights Reserved.

Muscles categorization
o Structure
• Striated: skeletal and cardiac muscles
• Unstriated: smooth muscles
o Control
• Voluntary: skeletal muscles
• Involuntary: cardiac and smooth muscles

(a) Skeletal muscle (b) Cardiac muscle (c) Smooth muscle
3 Muscle Types
Multiple nuclei in Intercalated disc Smooth

single cell muscle cell
Cell nucleus Cell nucleus
Skeletal muscle
cell (muscle fiber) (Cells separated
Cardiac muscle
for clarity)
cell
Classification: Striated muscle, Classification: Striated muscle, Classification: Unstriated

voluntary muscle involuntary muscle muscle, involuntary muscle
Characteristics Description: Bundles of long, Description: Interlinked network Description: Loose network of
thick, cylindrical, striated, of short, slender, cylindrical, short, slender, spindle- shaped,
contractile, multinucleate cells striated, branched, contractile unstriated, contractile cells that
that extend the length of the cells connected cell to cell by are arranged in sheets
muscle intercalated discs
Typical location: Walls of
Typical location: Attached to Location: Wall of heart hollow organs and tubes, such as
bones of skeleton stomach and blood vessels
Function: Movement of
Function: Movement of body in Function: Pumping of blood contents within hollow organs
relation to external environment out of heart

3 Muscle Types

Skeletal muscle
Cardiac muscle
for clarity)
cell


3 Muscle Types

Skeletal muscle
Cardiac muscle
for clarity)
cell


3 Muscle Types

Skeletal muscle
Cardiac muscle
for clarity)
cell


Skeletal Muscles
o Are composed of cells (fibers)

o Fibers are bound together by connective tissue
o Fibers communicate with nerves & blood vessels
o Contractions can play a part in:

• Homeostatic activities
 Heat generation
• Non-homeostatic activities
 Movement of the body through space

Skeletal Muscle Cells
o Are composed of myofibrils made of thick and

thin filaments
o Appear striated due to a highly organized

internal arrangement of thick and thin filaments:
sarcomeres

From the Skeletal Muscle to its Fiber

Skeletal Muscle Cells
Sarcolemma:
Muscle fiber plasma (cell) membrane
Sarcoplasm:
Muscle fiber cytoplasm
o Contains glycosomes (granules of glycogen)
o Contains the oxygen-binding protein called myoglobin
o Is almost completely filled with contractile filaments
called myofilaments

The Sarcoplasmic Reticulum (SR)
o It is the Smooth Endoplasmic Reticulum (SER)

in the muscle fiber
o It is a network of tubes surrounding myofibrils
o It releases calcium ions to allow contraction
o It reabsorbs calcium ions during relaxation

The Transverse Tubules (T Tubules)
o Formed by invaginations of the sarcolemma and

flanked by the sarcoplasmic reticulum
o Are continuous with the sarcolemma
o Provide tightly linked signals for muscle
contraction
o Conduct electrical impulses to the throughout
cell (every sarcomere)
• The electrical impulse signals for the release of Ca2+
from adjacent terminal cisternae
Sarcolemma & Sarcoplasm

The Triad structure (TT & SR)

The Triad structure (TT & SR)

Triad function: DHPR & RyR
o T tubule proteins are the Dihydropyridine receptors (DHPR)

Calcium channels
Act as voltage sensors
o SR foot proteins are Ryanodine receptors (RYR)
Calcium channels
Regulate Ca2+ release from the SR cisternae

The Sarcomere
A band
Thick filaments along with portions of thin filaments that overlap
I band
Remaining portion of thin filaments that do not project into A band
Cross-bridges
Project from each thick filament in six directions toward the surrounding thin
filaments
Myosin
Myosin forms thick filaments

o Protein consisting of two identical subunits, each
shaped somewhat like a golf club

Actin
Actin is the main component of thin filaments

o Interacts with the myosin cross-bridges
o Two other proteins, tropomyosin and troponin, lie
across the surface of actin

Tropomyosin and Troponin
Tropomyosin lies alongside groove of actin

spiral:
• Covers actin sites blocking cross-bridge binding
Troponin is made of three polypeptide units:

• One binds to tropomyosin
• One binds to actin
• One binds with Ca2+

Bands and Cross-Bridges
A and I bands
A: thick filaments along with portions of thin filaments that overlap
I: remaining portion of thin filaments that do not project into A band
Cross-bridges
Project from each thick filament in six directions toward the
surrounding thin filaments
Molecular Basis of the Contraction
During contraction
Cycles of cross-bridge binding and bending pull
thin filaments inward
o Sliding filaments mechanism
• Contraction is accomplished by thin filaments from the opposite
sides of each sarcomere sliding closer together between the
thick filaments
o Power stroke
• Stroking motion pulls the thin filament toward the center of the
sarcomere

Calcium (Ca2+)
The link Between Excitation and Contraction
Stimulation by the neuron (SNS)
ÞA graded potential is formed at the end plate
ÞThis stimulates Action Potentials to propagate across the sarcolemma
ÞThe DHP is activated by the Action Potentials
ÞActivated DHP interact with Ryanodine receptors in the lateral sacs of the SR
ÞThe Ryanodine receptors open to allow Ca 2+ to leak into the cytosol
ÞCa2+ binds with the troponin, initiating the contractile cycle

Calcium (Ca2+)
The link Between Excitation and Contraction

Fiber at rest
1 Energized: ATP split by myosin
ATPase; ADP and Pi remain attached to
myosin; energy stored in cross bridge ADP ADP
y
(that is, energy “cocks” cross bridge). ...or... No Ca2+
erg
erg
En
En
2b Resting: No excitation; no
Ca2+ released; actin and myosin
prevented from binding; no
cross-bridge cycle; muscle fiber
remains at rest.
present (excitation)
4a Detachment: Linkage
ADP
y
between actin and myosin Cross-
erg
2a Binding: Ca2+ released on
broken as fresh molecule of bridge
En
excitation; removes inhibitory
ATP binds to myosin cross cycle influence
bridge; cross bridge assumes
from actin, enabling it to
original
bind with cross bridge.
conformation; ATP hydrolyzed
(cycle starts again at step 1).
Fresh ATP available

ADP
Energy
3 Bending: Power stroke of

...or... cross bridge triggered on
contact between myosin and
actin; Pi released during and
No ATP (after death) ADP released after power
stroke.
4b Rigor complex: If
no fresh ATP available
(after death),
actin and myosin remain
bound in rigor complex.

Cross-bridge cycle
y
erg
erg
En
En
remains at rest.
ADP
y
erg
En
original
Fresh ATP available

ADP
Energy

stroke.
(after death),

Cross-bridge cycle
y
erg
erg
En
En
remains at rest.
ADP
y
erg
En
original
Fresh ATP available

ADP
Energy

stroke.
(after death),

Cross-bridge cycle
y
erg
erg
En
En
remains at rest.
ADP
y
erg
En
original
Fresh ATP available

ADP
Energy

stroke.
(after death),

Cross-bridge cycle
y
erg
erg
En
En
remains at rest.
ADP
y
erg
En
original
Fresh ATP available

ADP
Energy

stroke.
(after death),

Cross-bridge cycle
y
erg
erg
En
En
remains at rest.
ADP
y
erg
En
original
Fresh ATP available

ADP
Energy

stroke.
(after death),

Cross-bridge cycle
Skeletal muscle contraction is voluntary and controlled by the somatic nervous
system
1. A neural signal results in Ca2+ being released from the lateral sacs of the
sarcoplasmic reticulum
2. This Ca2+ binds with troponin, which alters the shape of the
troponin/tropomyosin complex unmasking the myosin binding sites on the
actin monomers
3. This results in the formation of the crossbridge where myosin binds with actin
4. Once the crossbridge has formed, the power stroke takes place with the
displacement of ATP and Pi from the myosin head
5. The power stroke progresses to form the rigor complex, a very strong bond
between the actin and myosin molecules
6. The power stroke causes the thin filaments to be pulled over the thick
filaments, shortening the sarcomere
7. In order to release the bond between actin and myosin, ATP must bind to the
myosin head and be hydrolyzed to ADP and Pi. This sequence of crossbridge
and power stroke continues until the sarcomere is fully contracted
Latent Contraction Relaxation
Relationship of an action potential period time time
to the resultant muscle twitch in a
Muscle
slow contracting fiber twitch
Tension
Contractile
response
A few
msec
+30
Membrane potential (mV)
Action
0 potential
Note that the resting potential –90

of a skeletal muscle fiber is –90 1 msec
mV, compared to a resting 50 100

potential of –70 mV in a neuron Stimulation Time (msec)

Skeletal Muscle Mechanics
o Tendons attach the muscle to bones
o Muscle tension is transmitted to bone as the

contractile component tightens the series-
elastic component (tendon)
• Tension is produced internally within the sarcomeres
 Contractile component of the muscle

Skeletal Muscles of the Human Body

Lever Systems
Interactive units of skeletal muscles, bones,

and joints form lever systems
o Lever: rigid structure capable of moving around a

pivot point known as a fulcrum
• Bones are levers

• Joints are fulcrums
• Skeletal muscles provide the force to move the bones

Flexion Extension
Origin Origin
of biceps of triceps
Biceps Triceps
contracts contracts
Insertion Insertion
of biceps of triceps

Primary Types of Contraction
Isotonic (constant tension):

o Load remains constant as the muscle changes
Isokinetic (constant motion):

o Velocity remains constant as the fibers shorten
Isometric (constant length):

o Muscle length remains constant as tension increases

Other Types of Contractions
Concentric contractions
o Muscle shortens
Eccentric contractions
o Muscle lengthens
PS: Some skeletal muscles do not attach to

bones at both ends but still produce
movement (e.g., tongue muscles)

Load, Velocity, Work, and Force
The velocity of shortening is related to the load

o Load–velocity relationship
Although muscles can accomplish work, much of

the energy is converted to heat
o Work: force multiplied by distance
o Force: muscle tension required to overcome
the load (the weight of the object)

Graded Contractions
Contractions of a whole muscle can be of

varying strength
Two primary factors can be adjusted to

produce graded contractions:
• Number of muscle fibers contracting

• Tension developed by each contracting fiber

Number of fibers
The number of fibers contracting within a

muscle depends on the extent of motor unit
recruitment
o More motor units are recruited for stronger

contractions

Motor neuron
Muscle fiber
Spinal cord
KEY
=
Motor unit 1
=
Motor unit 2
=
Twitch Summation
Frequency of stimulation can influence the tension

developed by each muscle fiber
Twitch summation and tetanus

o Twitch summation: the increase
in tension accompanying
repetitive stimulation of a
muscle fiber
o Tetanus: a sustained contraction
of maximal strength

Twitch Summation
Twitch summation results primarily from a sustained

elevation in cytosolic Ca2+
o Additional factor: more time to stretch the series-elastic
component

Muscle Metabolism & Fiber Types
Four steps in the excitation, contraction, and

relaxation processes require ATP
o Splitting of ATP by myosin ATPase provides the
energy for power stroke of cross bridge
o Binding of ATP to myosin breaks cross bridge
o Active transport of Ca2+ back into lateral sacs of the

SR during relaxation
o Activity of Na+-K+ pump during action potential
production

Alternate Pathways
Muscle fibers have alternate pathways for forming new ATP
o Creatine phosphate: is the first energy storehouse used at the onset of

contractile activity. Creatine kinase is the enzyme responsible for liberating the
energy needed for synthesizing ATP
o Oxidative phosphorylation: require adequate supply of oxygen so these
muscles are replete with myoglobin. These muscles do not fatigue easily.
o Glycolysis: utilized during high-intensity exercise, when oxygen may be
limited. These muscles produce powerful contractions but fatigue easily
o Lactate production: At times of intense exercise, the glycolytic process
may overwhelm oxidative phosphorylation, and the glycolytic end products
(pyruvate) may build up in the muscle. The muscle will convert the pyruvate to
lactate by the process of fermentation.

Fatigue
Fatigue may be of muscle or central origin

o Muscle fatigue occurs when an exercising muscle can
no longer respond to stimulation with the same degree of
contractile activity
The primary factors involved include:

– a local increase in inorganic phosphate, which reduces the strength
of the myosin/actin interaction and decreases the sensitivity of the
troponin/tropomyosin complex to Ca++.
– the depletion of glycogen stores plays a role in fatigue.

Fatigue

no longer respond to stimulation with the same
degree of contractile activity
o Central fatigue occurs when the CNS no longer
adequately activates motor neurons

Fatigue

no longer respond to stimulation with the same
degree of contractile activity
o Central fatigue occurs when the CNS no longer
adequately activates motor neurons
Increased O2 consumption is necessary to

recover from exercise
o Excess post-exercise oxygen consumption

Types of Skeletal Muscle Fibers
Based on differences in ATP hydrolysis and

synthesis
o Slow-oxidative (type I) fibers

o Fast-oxidative (type IIa) fibers
o Fast-glycolytic (type IIx) fibers

Comparing Skeletal Muscle Fibers
Fast versus slow fibers

o Fast fibers have higher myosin ATPase (ATP-splitting)
activity than slow fibers
Oxidative versus glycolytic fibers

o Fiber types differ in ATP-synthesizing ability
Genetic endowment of muscle fiber types

o Largely determined by the type of activity for which
the muscle is specialized

Muscle fiber types
Fast twitch
Tension (% of maximum)
Slow twitch
Comparison of the speed of contraction
of fast and slow muscle fiber types
0 20 40 60 80 100
Time
Stimulation (msec)
(a)
Cross section of a human

muscle showing the distribution
muscle fiber types
Slow-oxidative fiber Fast-oxidative fiber Fast-glycolytic fiber
(b)
Muscle Fibers and Adaptation
Muscle fibers adapt considerably in response

to the demands placed on them
o Improvement in oxidative capacity

o Muscle hypertrophy
o Influence of testosterone
o Interconversion between fast muscle types
o Muscle atrophy
o Limited repair of muscle

Muscle Fibers and Adaptation
• Regular aerobic exercise will promote metabolic changes within oxidative muscle
fibers, allowing them to more efficiently use O2 and therefore better tolerate
prolonged exercise without fatiguing.
• Muscle fibers can also respond to regular bouts of anaerobic, short-duration, high-
intensity resistance training by undergoing hypertrophy  an enlargement of the
muscle due to increased synthesis of actin and myosin filaments.
• Muscles may atrophy (decrease mass) due to:

(1) disuse atrophy, seen in muscles not used for long periods of time, such as when a
limb is in a cast;
(2) denervation atrophy, if the nervous input to the muscle is lost
(3) age-related atrophy or sarcopenia, which begins at about age 40 with progressive
loss of motor neurons, especially those innervating fast-glycolytic fibers.

Control of Motor Movement
Motor activity can be classified as reflex,

voluntary, or rhythmic
o Somatic reflex responses
o Voluntary movements
o Rhythmic activities
Multiple neural inputs influence motor unit

output
o Input from afferent neurons, the primary motor cortex,
and brain stem

Muscle Receptors
Muscle receptors provide afferent information

needed to control skeletal muscle activity
o For effective control of motor output, the CNS needs

continual information regarding ongoing changes in
muscle length and tension
• Muscle spindles monitor muscle length

• Golgi tendon organs detect changes in tension

Muscle Spindles and the Stretch Reflex
Muscle spindle structure

o Distributed throughout the fleshy part of a skeletal
muscle
o Consist of collections of specialized muscle fibers
known as intrafusal fibers
Stretch reflex
o Local negative-feedback mechanism to sense and
resist changes in muscle length when an additional
load is applied

Motor Neurons & Golgi Tendon Organs
Coactivation of gamma and alpha motor

neurons
o Gamma motor neurons initiate contraction of the
muscular end regions of intrafusal fibers
o Alpha-gamma coactivation: simultaneous stimulation
of gamma and alpha motor-neuron system
Golgi tendon organs

o Located in muscle tendons
o Respond to changes in the muscle’s tension

Pain Reflexes
Skeletal muscle reflexes can be triggered by

painful stimulation of the skin
o Withdrawal reflex: used to withdraw from a painful

stimulus
o Crossed extensor reflex: ensures the opposite limb is

in a position to bear the weight of the body as the
injured limb is withdrawn from the stimulus

Points to Ponder
o What makes one muscle larger than

another muscle?
o What causes your foot to “fall asleep”?
o Why do you think the average life

expectancy for NFL players is only 57
years?

Muscle Physiology

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Human Physiology – From Cells to Systems | 9e

Prof. Walid Hleihel

o Largest group of tissues in the body

o Constitute approximately half the body’s weight

• Skeletal muscles accounting for

 40% of the body weight in men

© Cengage Learning 2016. All Rights Reserved.

• Striated: skeletal and cardiac muscles

• Unstriated: smooth muscles

• Voluntary: skeletal muscles

• Involuntary: cardiac and smooth muscles

© Cengage Learning 2016. All Rights Reserved.

Multiple nuclei in Intercalated disc Smooth

Classification: Striated muscle, Classification: Striated muscle, Classification: Unstriated

© Cengage Learning 2016. All Rights Reserved.

Multiple nuclei in Intercalated disc Smooth

Classification: Striated muscle, Classification: Striated muscle, Classification: Unstriated

© Cengage Learning 2016. All Rights Reserved.

Multiple nuclei in Intercalated disc Smooth

Classification: Striated muscle, Classification: Striated muscle, Classification: Unstriated

© Cengage Learning 2016. All Rights Reserved.

Multiple nuclei in Intercalated disc Smooth

Classification: Striated muscle, Classification: Striated muscle, Classification: Unstriated

© Cengage Learning 2016. All Rights Reserved.

o Are composed of cells (fibers)

o Contractions can play a part in:

© Cengage Learning 2016. All Rights Reserved.

o Are composed of myofibrils made of thick and

o Appear striated due to a highly organized

© Cengage Learning 2016. All Rights Reserved.

© Cengage Learning 2016. All Rights Reserved.

© Cengage Learning 2016. All Rights Reserved.

o It is the Smooth Endoplasmic Reticulum (SER)

© Cengage Learning 2016. All Rights Reserved.

o Formed by invaginations of the sarcolemma and

© Cengage Learning 2016. All Rights Reserved.

© Cengage Learning 2016. All Rights Reserved.

© Cengage Learning 2016. All Rights Reserved.

o T tubule proteins are the Dihydropyridine receptors (DHPR)

© Cengage Learning 2016. All Rights Reserved.

Myosin forms thick filaments

© Cengage Learning 2016. All Rights Reserved.

Actin is the main component of thin filaments

© Cengage Learning 2016. All Rights Reserved.

Tropomyosin lies alongside groove of actin

Troponin is made of three polypeptide units:

© Cengage Learning 2016. All Rights Reserved.

© Cengage Learning 2016. All Rights Reserved.

© Cengage Learning 2016. All Rights Reserved.

© Cengage Learning 2016. All Rights Reserved.

Fresh ATP available

3 Bending: Power stroke of

© Cengage Learning 2016. All Rights Reserved.

Fresh ATP available

3 Bending: Power stroke of

© Cengage Learning 2016. All Rights Reserved.

Fresh ATP available

3 Bending: Power stroke of

© Cengage Learning 2016. All Rights Reserved.

Fresh ATP available

3 Bending: Power stroke of

© Cengage Learning 2016. All Rights Reserved.

Fresh ATP available