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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
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Muscle contraction
• The plasma membrane of muscle cells or fibers is called the sarcolemma.
•
The cytoplasm of muscle cells is called sarcoplasm.
• Within the sarcoplasm, bundles of myofibrils are arranged in parallel
• Myofibrils are made up of bundles of protein myofilaments organized in a basic pattern that is repeated in the
space

• We distinguish between thin actin myofilaments (with tropomyosin and troponin as regulatory elements) and myofilaments
thick myosin
• Myofibrils are surrounded by the sarcoplasmic reticulum.
• The transverse tubules (T) are invaginations of the sarcolemma that are arranged parallel to the terminal cisternae of the
sarcoplasmic reticulum

MUSCLE FIBER MYOFIBRILS MYOFILAMENTS

(cell) (bundles of myofilaments) (actin/myosin)
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Muscle contraction
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Muscle contraction

Thin filament
A thin filament consists of two strands of actin subunits twisted into a
helix plus two types of regulatory proteins (troponin and tropomyosin).

Portion of a thin filament

Tropomyosin Troponin Actin

Active sites
for myosin
Actin subunits attachment

Actin subunits
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Muscle contraction

thick filament
Each thick filament consists of many myosin molecules whose heads
protrude at opposite ends of the filament.

Portion of a thick filament

Myosin head

Actin-binding sites

Heads Tail
ATP-
binding
site
Flexible hinge region
Myosin molecule
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Muscle contraction
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Muscle contraction
Z M Z

H-zone

I-band A-band I-band

sarcomere
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Muscle contraction
Sliding filament model of contraction.

Thin filament (actin) Myosin heads Thick filament (myosin)

Z Z
h
TO
Transmission electron micrograph of part of a sarcomere clearly Yo Yo

showing the myosin heads forming cross bridges that generate

the contractile force. 1 Fully relaxed sarcomere of a muscle fiber

Z Z

muscle contraction animation Yo TO Yo

2 Fully contracted sarcomere of a muscle fiber

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Muscle contraction
Types of skeletal muscle fibers

• Fibers I (red, slow oxidative)

•Smaller diameter
•Slower contraction

•Abundant myoglobin, more reddish color

• IIA fibers (intermediate, fast oxidative)

•Similar to white

•Greater resistance to fatigue

• IIX fibers (white, fast glycolytic)

•Whitish color (ATPase)
•Larger diameter
•Abundant glycogen
•Densely packed myofibrils
•Relatively few mitochondria
•Little myoglobin, pale color
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Muscle contraction
Types of skeletal muscle fibers

• Fibers I (red, slow oxidative)

•Smaller diameter
•Slower contraction

•Abundant myoglobin, more reddish color

• IIA fibers (intermediate, fast oxidative)

•Similar to white

•Greater resistance to fatigue

• IIX fibers (white, fast glycolytic)

•Whitish color (ATPase)
•Larger diameter
•Abundant glycogen
•Densely packed myofibrils
•Relatively few mitochondria
•Little myoglobin, pale color
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Muscle contraction
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Muscle contraction
(Type I) (IIA) (IIX)
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Muscle contraction

(a) Direct phosphorylation (b) Anaerobic pathway (c) Aerobic pathway

Coupled reaction of creatine phosphate Glycolysis and lactic acid formation Aerobic cellular respiration
(CP) and ADP

Energy source: CP Energy source: glucose Energy source: glucose; pyruvic acid; free
fatty acids from adipose tissue; amino
acids from protein catabolism

CP ADP Glucose (from Glucose (from

glycogen breakdown glycogen breakdown
Creatine or delivered from blood) or delivered from blood)
kinase O2
Glycolysis Pyruvic acid
Creatine ATP in cytosol Fatty
acids O2
O2
2 ATP amino Aerobic respiration
Aerobic respiration
in mitochondria
in mitochondria
Pyruvic acid acids
net gain
O2 32 ATP
Released Lactic acid CO2
to blood H2O net gain per
glucose

Oxygen use: None Oxygen use: None Oxygen use: Required

Products: 1 ATP per CP, creatine Products: 2 ATP per glucose, lactic acid Products: 32 ATP per glucose, CO2, H2O
Duration of energy provision: 15 Duration of energy provision: 60 Duration of energy provision: Hours
seconds seconds, or slightly more
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Muscle contraction

Predominance Small load Predominance

of fast glycolytic of slow oxidative
(fatigable) fibers (fatigue-resistant)
fibers

Contractile Contractile
velocity duration
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Muscle contraction

Prolonged-duration
Short-duration exercise
exercise

ATP stored in ATP is formed Glycogen stored in muscles is broken ATP is generated by
muscles from creatine down to glucose, which is oxidized to breakdown of several
is used first. Phosphate generate ATP. nutrient energy fuels by
and ADP. aerobic pathway. This
pathway uses oxygen
released from myoglobin
or delivered in the blood
by hemoglobin. When it
ends, the oxygen deficit is
paid back.
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Muscle contraction

Spinal cord
Axon terminals at
Branching axon to
neuromuscular junctions motor unit
Engine Engine
unit 1 unit 2

Nerve
motor neuron cell
body
motor neuron
axon

muscle

Muscle
fibers

(b) Branching axon

terminals form
neuromuscular
Axons of motor neurons extend from the spinal cord to the muscle. (to)
There each axon divides into a number of axon terminals that form neuromuscular junctions, one per
junctions with muscle fibers scattered throughout the muscle. muscle fiber (photo-
micrograph 330x).
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Muscle contraction
Action potential (AP) arrives at
axon terminal at neuromuscular junction

ACh released; binds to receptors

on sarcolemma

Phase 1 Ion permeability of sarcolemma changes

Muscle fiber
is stimulated by motor
neuron.
Local change in membrane
voltage (depolarization) occurs

Local depolarization (end

The phases leading to muscle fiber plate potential) ignites AP in sarcolemma
contraction.

AP travels across the entire sarcolemma

AP travels along T tubule

Phase 2 SR releases Ca2+; Ca2+ binds

Excitation-contraction to troponin; myosin-binding
coupling occurs. sites (active sites) on actin exposed

Myosin heads bind to

actin; contraction begins
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Muscle contraction
Action Myelinated axon
potential (AP) of motor neuron
Axon terminal
Nucleus of neuromuscular
junction
Sarcolemma
of the muscle fiber

1 Action potential arrives at

axon terminal of motor neuron.
Ca2+ Synaptic vesicle
Ca2+
2 Voltage-gated Ca2+ channels containing ACh
open and Ca2+ enters the Mitochondrion
axon terminal. Axon terminal of Synaptic
motor neuron cleft
3 Ca2+ entry causes some Fusing synaptic
vesicles
synaptic vesicles to release
their contents (acetylcholine) Junctional
by exocytosis. ACh
folds of
sarcolemma
4 Acetylcholine, a
neurotransmitter, diffuses across Sarcoplasm of
the synaptic cleft and binds muscle fiber
to receptors in the sarcolemma.
5 ACh binding opens ion Na+ K+ Postsynaptic membrane
channels that allow simultaneous ion channel opens;
passage of Na+ into the muscle ions pass.
fiber and K+ out of the muscle
fiber.
6 ACh effects are terminated Ach– Degraded ACh
Na+ Postsynaptic membrane
by its enzymatic breakdown ion channel closed; Nicotinic ACh receptor
in the synaptic cleft ions cannot pass.
by acetylcholinesterase. Acetyl-
cholinesterase
K+
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Muscle contraction
Na+ channels
close, K+ channels
Depolarization due open
to Na+ entry

Repolarization
due to K+ exit
Na+
channels
open Threshold

terminal axon K+ channels

close

Open Na+ Closed K+

channel channel
Na+
Synaptic
cleft

ACh–
Na+ K+ K+
ACh
Na+ K+
2 Generation and propagation of the action
potential (AP)
Closed Na+ Open K+
channel channel
1 Local depolarization: Na+
generation of the end plate
potential on the sarcolemma

K+
Sarcoplasm of muscle fiber 3 Repolarization

Figure 9.9 Summary of events in the generation and propagation of an action potential in a skeletal muscle fiber.
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Muscle contraction
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Muscle contraction
During the EC coupling of skeletal muscle, acetylcholine receptors in the
sarcolemmal (plasma) membrane of skeletal muscle cells are activated by
acetylcholine released from a motor neuron.

Acetylcholine receptors are ligand-gated Na+ channels, through which Na+

ions rush into the cytosol of skeletal muscle fibers. The Na+ influx induces
the depolarization of the sarcolemmal membrane in skeletal muscle fibers.

The membrane depolarization spreading along the surface of the sarcolemmal

membrane reaches the interior of skeletal muscle fibers via the invagination
of the sarcolemmal membranes (t-tubules).

Dihydropyridine receptors (DHPRs, a modified voltage-gated Ca2+ channel

on the t-tubule membrane) are activated by the depolarization of the t- tubule
membrane, which in turn activates ryanodine receptor 1 (RyR1, a ligand-
ACh-induced depolarization gated Ca2+ channel on the sarcoplasmic reticulum (SR) membrane) via
physical interaction.

DHPR DHPR
Ca2+ ions that are stored in the SR are released to the cytosol via the
activated RyR1, where they bind to troponin C, which then activates a series
Plasma of contractile proteins and induces skeletal muscle contraction.
membrane
It is worth noting that during skeletal EC coupling, the contraction of skeletal
muscle occurs even in the absence of extracellular Ca2+ because DHPR
mechanical coupling serves as a ligand for RyR1 activation via physical interactions. The Ca2+
Cytosol
entry via DHPR is not a necessary factor for the initiation of skeletal muscle
contraction, although Ca2+ entry via DHPR does exist during skeletal EC
coupling.
MR
membrane

RyR1 RyR1

Ca2+
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Muscle contraction
Steps in EC Coupling:
Sarcolemma
Voltage-sensitive T tube 1 Action potential is propagated along
tubular protein
the sarcolemma and down the T tubules.

Ca2+
release
channel
2 Calcium ions are released.
Tanker
terminal
of SR

Ca2+

Actin

Tropomyosin
Ca2+ Troponin blocking active sites
Myosin

Calcium binds to troponin and

3 removes the blocking action of
tropomyosin.

Active sites exposed and

ready for myosin binding

4 Contraction begins

Myosin
cross
bridge

The aftermath
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Muscle contraction
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Muscle contraction

Ca +2 ACTS AS A REGULATOR OF CONTRACTION

• AT REST (LOW INTRACELLULAR Ca+2 ) THE C SUBUNIT OF TROPONIN (TnC) IS RELAXED
• ACTIN-MYSIN CROSS BRIDGES CANNOT BE ESTABLISHED BECAUSE TROPOMYOSIN IS INTERPENTED BETWEEN THE BOTH
• THE INCREASE IN INTRACELLULAR Ca+2 CAUSES TnC TO CHANGE CONFORMATION.
• THIS CONFORMAL CHANGE DISPLACES TROPOMIOSIN
• MYOSIN HEADS CAN NOW INTERACT WITH ACTIN
• THE MYOFILAMENTS SLIDE, SHORTENING THE SARCOMERES (CONTRACTION)
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Muscle contraction

45º

45º

~11nm

90º

~11nm
90º

https://derangedphysiology.com/main/cicm-primary-exam/required-reading/musculoskeletal-
system/Chapter%20131/mechanism-excitation-contraction-coupling

45º
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Muscle contraction
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Muscle contraction

Isotonic

Two (3) types of contraction

Isotonic contraction: Muscle contracts at constant tension

• Concentric: muscle shorthens during contraction •

Eccentric: muscle elongates during contraction

Isometric contraction: Muscle contracts at constant length

Isometric
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Muscle contraction

(a) Concentric isotonic contraction (a) Concentric isotonic contraction

On stimulation, muscle develops enough tension (force) to lift the load
(weight). Once the resistance is overcome, the muscle shortens, and the
tension remains constant for the rest of the contraction. Amount Muscle
of resistance relaxes

Peak tension
developed
Tendon

muscle
stimulus
Muscle Resting length
contracts
(isotonic
contraction) 3kg
Tendon

3kg
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Muscle contraction

light load

Intermediate load

Heavy load

Stimulus
(a) The greater the load, the less the muscle shortens (b) The greater the load, the slower
and the shorter the duration of contraction the contraction
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Muscle contraction

(b) Isometric contraction (b) Isometric contraction

Muscle is attached to a weight that exceeds the muscle's peak tension-
developing capabilities. When stimulated, the tension increases to the
muscle's peak tension-developing capability, but the muscle does not
shorten. Amount of resistance
Muscle
relaxes
Peak tension
developed

muscle
stimulus
Muscle Resting length
contracts
(isometric
contraction)

6kg 6kg
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Muscle contraction

Hill's model of muscle

• One contractile element (CE) (sarcomeres)

• Two non-linear spring elements, in series (SE) and in parallel (PE)

• The parallel element represents the connective tissue that surrounds the
contractile element (fascia, epi-peri- and endomysium)

• The parallel element is responsible for the muscle passive behavior

when it is stretched

• The series element represents the tendon and the intrinsic elasticity of
the myofilaments (titin...)
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Muscle contraction

Length-tension relationship
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Muscle contraction
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Muscle contraction

Single stimulus single twitch

contraction
relaxation

Stimulus

(a) A single stimulus is delivered. The muscles

contracts and relaxes
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Muscle contraction

Latent Period of Period of

period contraction relaxation

Single
stimulus
(a) Myogram showing the three phases of an isometric twitch
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Muscle contraction

Latent period

Extraocular muscle (lateral rectus)

Gastrocnemius
Soleus

Single
stimulus
(b) Comparison of the relative duration of twitch responses of
three muscles
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Muscle contraction

Large Muscle and

number of sarcomere
Large High stretched to
muscle frequency of slightly over 100%
activated musclefibers
fibers stimulation of resting length

Contractile force
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Muscle contraction
Single stimulus single twitch

contraction
relaxation

Stimulus

(a) A single stimulus is delivered. The muscles

contracts and relaxes

Low stimulation frequency High stimulation frequency fused

unfused (incomplete) tetanus (complete) tetanus

Partial relaxation

Stimuli Stimuli

(b) If another stimulus is applied before the muscle relaxes

completely, then more tension results. (c) At higher stimulus frequencies, there is no relaxation
This is temporary (or wave) summation and results in unfused at all between stimuli. This is fused (complete) tetanus.
(or incomplete) tetanus.

Temporary summation
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ABSOLUTE PR
RELATIVE PR

ABSOLUTE PR
RELATIVE PR

SHORT REFRACTORY PERIOD:

TETANIZATION IN SKELETAL Ms.

IN THE SKELETAL MUSCLE

TETANIZATION POSSIBLE
STRONG AND SUSTAINED CONTRACTION
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Muscle contraction

Stimulus strength

Maximal
stimulus

Stimulus
Threshold

Proportion of motor units excited

Strength of muscle contraction

Maximal contraction

Spatial summation: recruitmenof motor units

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Muscle contraction

Engine Motor Motor

unit 1 unit 2 unit 3
Recruited recruited recruited
(small (medium (large
fibers) fibers) fibers)

Spatial summation: recruitmenof motor units

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Muscle contraction

• A cross section of a muscle 5 cm in diameter

It contains 800 x 109 thick myosin filaments.

• Each thick filament contains ~300 myosin heads.

• That 5 cm diameter section of muscle contains 2 x

1014 myosin heads working in parallel.

•
2 x 1014 is like the total population of the earth multiplied by
10,000

• Assuming that this muscle is capable of exercising a

force of 225 N, we would have that each myosin head
develops a force of around 1 pN.

• By carrying out direct measurements with an optical tweezer,

proven that a single myosin molecule can generate a
movement of ~ 10 nm against a force of ~ 3 pN
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Muscle contraction

Types of smooth muscle

Unitary (visceral)

• Gap junctions between adjacent cells

• Electrical syncytium

• Myogenic contraction (pacemaker) Longitudinal layer of

smooth muscle (shows
• Modular by SNA smooth muscle fibers in
cross section)
• The most abundant:
• digestive tract •
blood vessels (except elastic arteries) • urinary
tract...

Multiunit small intestine

Mucous membrane

• No gap junctions (b) Cross section of the

Circular layer of smooth
intestine showing the
• Neurogenic contraction muscle (shows longitudinal
smooth muscle layers (one
views of smooth muscle fibers)
• Fundamentally in: • circular and the other longitudinal)
trachea running at right angles to each other.
• iris
• elastic arteries
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Muscle contraction
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Muscle contraction
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Muscle contraction

Intermediate Caveolae Gap junctions

filament

Nucleus Dense bodies

(a) Relaxed smooth muscle fiber (note that adjacent fibers
are connected by gap junctions)

Nucleus
Dense bodies

(b) Contracted smooth muscle fiber

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Muscle contraction
Extracellular fluid (ECF)
Ca2+
Plasma membrane

Cytoplasm

1 Calcium ions (Ca2+)

enter the cytosol from
the ECF via voltage- Extracellular fluid (ECF)
dependent or voltage- Ca2+
independent Ca2+
channels, or from
Ca2+
the scant SR. Plasma membrane
2 Sarcoplasmic
Ca2+ binds to and
reticulum
activates calmodulin.

Ca2+

Inactive calmodulin Activated calmodulin

Cytoplasm
3 Activated calmodulin
activates the myosin
light chain kinase 1
enzymes.
Inactive kinase Activated kinase Calcium ions
4
ATP
(Ca2+) enter the
The activated kinase enzymes
catalyze transfer of phosphate to
ADP
cytosol from the
myosin, activating the myosin
ATPases. ECF via voltage- Ca2+
Pi
dependent or
Pi
voltage-
Inactive
myosin molecule
Activated (phosphorylated)
myosin molecule
independent Ca2+ channels, or from the scant SR.
5 Activated myosin forms cross
bridges with actin of the thin
filaments and shortening begins. Sarcoplasmic
reticulum
Thin
filament

thick
filament

Sequence of events in excitation-contraction coupling of smooth muscle.

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Muscle contraction
Extracellular fluid (ECF)
Ca2+
Plasma membrane

Cytoplasm

1 Calcium ions (Ca2+)

enter the cytosol from
the ECF via voltage-
dependent or voltage- Ca2+
independent Ca2+
channels, or from
the scant SR.

2 Sarcoplasmic
Ca2+ binds to and
reticulum
activates calmodulin.

Ca2+

Inactive calmodulin Activated calmodulin 2 Ca2+ binds to and

activates calmodulin.
3 Activated calmodulin
activates the myosin
light chain kinase
enzymes.
Inactive kinase Activated kinase Ca2+
ATP
4 The activated kinase enzymes
catalyze transfer of phosphate to
myosin, activating the myosin ADP
ATPases.
Inactive calmodulin Activated calmodulin
Pi
Pi

Inactive Activated (phosphorylated)

myosin molecule myosin molecule

5 Activated myosin forms cross

bridges with actin of the thin
filaments and shortening begins.

Thin
filament

thick
filament

Sequence of events in excitation-contraction coupling of smooth muscle.

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Muscle contraction
Extracellular fluid (ECF)
Ca2+
Plasma membrane

Cytoplasm

1 Calcium ions (Ca2+)

enter the cytosol from
the ECF via voltage-
dependent or voltage- Ca2+
independent Ca2+
channels, or from
the scant SR.

2 Sarcoplasmic
Ca2+ binds to and
reticulum
activates calmodulin.

Ca2+

Inactive calmodulin Activated calmodulin

3 Activated calmodulin
3 Activated calmodulin
activates the myosin
activates the myosin
light chain kinase
light chain kinase
enzymes.
Inactive kinase Activated kinase enzymes.
4
ATP
Inactive kinase
The activated kinase enzymes
catalyze transfer of phosphate to
myosin, activating the myosin ADP
ATPases.

Pi
Pi

Inactive Activated (phosphorylated)

myosin molecule myosin molecule

5 Activated myosin forms cross

bridges with actin of the thin
filaments and shortening begins.

Thin
filament

thick
filament

Sequence of events in excitation-contraction coupling of smooth muscle.

Machine Translated by Google

Muscle contraction
Extracellular fluid (ECF)
Ca2+
Plasma membrane

Cytoplasm

1 Calcium ions (Ca2+)

enter the cytosol from
the ECF via voltage-
dependent or voltage- Ca2+
independent Ca2+
channels, or from
the scant SR.

2 Ca2+ binds to and

Sarcoplasmic
reticulum 4 ATP
activates calmodulin.
The activated kinase enzymes
Ca2+
catalyze transfer of phosphate
Inactive calmodulin Activated calmodulin
to myosin, activating the myosin ADP
3 Activated calmodulin
ATPases.
activates the myosin
light chain kinase
enzymes.
Inactive kinase Activated kinase

ATP
Pi
4 The activated kinase enzymes
catalyze transfer of phosphate to
myosin, activating the myosin ADP Pi
ATPases.

Pi
Pi Activated (phosphorylated)
Inactive
myosin molecule
Activated (phosphorylated)
myosin molecule
Inactivate myosin molecule
myosin molecule
5 Activated myosin forms cross
bridges with actin of the thin
filaments and shortening begins.

Thin
filament

thick
filament

Sequence of events in excitation-contraction coupling of smooth muscle.

Machine Translated by Google

Muscle contraction
Extracellular fluid (ECF)
Ca2+
Plasma membrane

Cytoplasm

1 Calcium ions (Ca2+)

enter the cytosol from
the ECF via voltage-
dependent or voltage- Ca2+
independent Ca2+
channels, or from
the scant SR.

Sarcoplasmic
5 Activated myosin forms
2 Ca2+ binds to and
activates calmodulin.
reticulum cross bridges with actin
Ca2+ of the thin filaments and shortening begins.
Inactive calmodulin Activated calmodulin

3 Activated calmodulin
activates the myosin
light chain kinase Thin
enzymes.
Inactive kinase Activated kinase
filament
ATP
4 The activated kinase enzymes
catalyze transfer of phosphate to
myosin, activating the myosin ADP
ATPases.

thick
Pi
Pi filament
Inactive Activated (phosphorylated)
myosin molecule myosin molecule

5 Activated myosin forms cross

bridges with actin of the thin
filaments and shortening begins.

Thin
filament

thick
filament

Sequence of events in excitation-contraction coupling of smooth muscle.

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Muscle contraction
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Muscle contraction

Single stimulus single twitch

contraction
relaxation

Stimulus

(a) A single stimulus is delivered. The muscles

contracts and relaxes
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Muscle contraction

Low stimulation frequency

unfused (incomplete) tetanus

Partial relaxation

Stimuli

(b) If another stimulus is applied before the muscle relaxes

completely, then more tension results.
This is temporary (or wave) summation and results in unfused
(or incomplete) tetanus.

Temporal summation: unfused tetanus

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Muscle contraction

High stimulation frequency fused

(complete) tetanus

Stimuli

(c) At higher stimulus frequencies, there is no relaxation

at all between stimuli. This is fused (complete) tetanus.

Temporal summation: fused tetanus

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Muscle contraction