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CHAPTER 44

LECTURE
SLIDES
Prepared by

Brenda Leady
University of Toledo
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Muscle
Vertebrates have three types of muscle
that are classified according to their
structure, function, and control
mechanisms
1. Cardiac muscle
2. Smooth muscle
3. Skeletal muscle

Skeletal muscle structure


Muscle is a grouping of cells (muscle
fibers) bound together by connective
tissue
Tendons link bones to skeletal muscle
Skeletal muscle fibers increase in size
during growth but few new fibers are
formed

Myofibrils

Striated muscle named for striped pattern

Myofibrils cylindrical bundles in muscle


fibers

One unit of repeating pattern is a


sarcomere

Thick filaments made of myosin

Thin filaments made of actin and other


proteins

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One muscle
fiber (cell)
One fascicle
(bundle of
muscle fibers)
Connective tissue

Muscle
Myofibril

Tendons

Blood vessels
and nerves

A band wide band of myosin


zone narrow region in center of A band, space
between the 2 sets of thin filaments

line in center of H zone, proteins that link central


regions of adjacent thick filaments

Z line Two sets of thin filaments anchored to


network of proteins at this point
Two

successive Z lines make a sarcomere

I band contains portions of thin filaments that


do not overlap thick filaments

Cross-bridges extend from surface of myosin


toward thin filaments

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Muscle fiber

Myofibril

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Muscle fiber

Myofibril

I band

A band

Myofibril

Z line

Z line
Sarcomere
8

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Muscle fiber

Myofibril

I band

A band

Myofibril

Z line

Z line
Sarcomere
M line

Z line

Z line

H zone
I band

A band

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Muscle fiber

Myofibril

I band

A band

Myofibril

Z line

Z line
Sarcomere
M line

Z line

Z line

H zone
I band

A band
Thin
filament

Actin

10

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Sarcomere
M line

Z line

Z line

H zone
I band

A band
Thin
filament

Thick (myosin) filament

Actin

Cross-bridge

11

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Muscle fiber

Myofibril

I band

A band

Myofibril

Z line

Z line
Sarcomere
M line

Z line

Z line

H zone
I band

A band
Thin
Actin
filament

Thick (myosin) filament

Cross-bridge

12

Sliding filament mechanism

Sarcomeres shorten as thin filaments slide


past stationary thick filaments

Myosin cross-bridges attach to thin filament


and force thin filament toward center of
sarcomere

Cross-bridge repeats motion as long as


stimulation to contract continues
13

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Relaxed

Shortened

Z line

A band

I band

A band
I band
unchanged reduced

H zone

M line

H zone
reduced

14

Actin
Each

actin molecule contains a binding site for


myosin

Molecules

form two intertwined helical chains

are closely associated with two proteins


tropomyosin and troponin that play important
roles in regulating contraction

Chains

15

Myosin
Six

protein subunits combine to form a protein with


two heads and a long tail

Tail

lies along axis of thick filament

Two

heads form cross-bridges

Each

head contains bonding site for actin and ATP

Myosin

proteins at 2 ends of thick filament oriented in


opposite directions so thin filaments are brought
toward center of sarcomere
16

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Thin filament

Actin

Tropomyosin

Troponin
Cross-bridge
Thick filament

Actin-binding site
ATP-binding site

Myosin tails

Cross-bridge
(2 hinges and 2 heads)

(a) Structure of thin and thick filaments


17

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Thin
filament

Thick
filament

H zone

Z line
Relaxed
M line

Shortened

H zone
Successive movements of
cross-bridges pull actin
toward the H zone.

(b) Cross-bridge mechanism


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Cross-bridge cycle

Events from the time when the cross-bridge


binds to a thin filament and when it is set to
repeat the process

Four steps
1.

Cross-bridge binds to actin

2.

Cross-bridge moves and filament slides past


each other power stroke

3.

ATP binds to myosin, and cross-bridge detaches

4.

Hydrolysis of ATP re-energizes the cross-bridge


19

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Thin filament (actin)

Binding:
When Ca2+ levels
are high, energized
cross-bridge can bind
to actin. (ADP + Pi are
already bound to the
cross-bridge.)

Ca2+

Z line
Thick filament
(myosin)

20

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Pi
2

Power stroke:
Release of Pi causes
cross-bridge to move
toward the H zone of
the sarcomere. This
power stroke moves
the actin filament
toward the H zone.
ADP is then released.

ADP

ATP

21

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Detaching:
ATP binds to myosin,
causing the crossBridge to detach from
the actin filament.

22

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Resetting:
Hydrolysis of ATP
to ADP+Pi provides
energy, which causes
the cross-bridge to
move away from the
H zone. ADP and Pi
remain bound to the
cross-bridge. Cycle
can begin again.

Energized
cross-bridge

23

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Thin filament (actin)


1

Binding:
When Ca2+ levels
are high, energized
cross-bridge can bind
to actin. (ADP + Pi are
already bound to the
cross-bridge.)

Ca2+

Z line
Thick filament
(myosin)

Pi
2

ADP

Power stroke:
Release of Pi causes
cross-bridge to move
toward the H zone of
the sarcomere. This
power stroke moves
the actin filament
toward the H zone.
ADP is then released.

ATP

Detaching:
ATP binds to myosin,
causing the crossBridge to detach from
the actin filament.

Resetting:
Hydrolysis of ATP
to ADP+Pi provides
energy, which causes
the cross-bridge to
move away from the
H zone. ADP and Pi
remain bound to the
cross-bridge. Cycle
can begin again.

Energized
cross-bridge

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Regulation of contraction

Tropomyosin
Rod-shaped

molecule composed of 2
intertwined proteins
Arranged along length of actin thin filament
In absence of calcium, cover myosin-binding
sites

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Troponin
Smaller

protein bound to both tropomyosin


and actin
Binds Ca2+ and drags tropomyosin off of
myosin-binding site and contraction begins
Removal of Ca2+ reverses the process and
contraction stops

26

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Tropomyosin

Troponin

Actin

Myosin-binding
sites are hidden.

Actin-bindingsite
Energized cross-bridge
cannot bind to actin.

(a) Low cytosolic Ca2+, relaxed muscle

Ca2+ binds to troponin,


causing tropomyosin to
move away from the
myosin-binding sites.

Ca2+

Myosin-binding sites
are exposed.

Cross-bridge
binds to actin
and generates
force.

(b) High cytosolic Ca2+, activated muscle

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Excitation-contraction coupling

Skeletal muscle cells are capable of generating and


propagating action potentials

Causes rise in cytosolic Ca2+ released from


sarcoplasmic reticulum

Transverse or T-tubules are invaginations of plasma membrane


that conduct the action potential from the outer surface to inside

Triggers contraction

Ion pumps will return calcium to the sarcoplasmic


reticulum, troponin and tropomyosin slide back in place,
and contraction stops
28

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1
Opening of transverse tubule
to extracellular fluid

Muscle fiber plasma


membrane

T-tubule

Action potentials propagate


along the plasma membrane
and down the transverse
tubules (T-tubules).

Muscle fiber
Sarcoplasmic
Plasma membrane reticulum

T-tubule

The depolarization produced by


the action potentials opens Ca2+
channels in the membranes of
the sarcoplasmic reticulum,
through which Ca2+ diffuses
into the cell cytosol.

Ca2+

Ca2+

ATP

Ca2+ totroponin
Ca2+
channel

ADP + Pi

Cytosol

Myofibrils
3

Cytosol

Ca2+ then binds


to troponin in
the myofibril,
initiating muscle
contraction.

Ca2+ is then pumped


back into the
sarcoplasmic
reticulum by ATP
dependent ion pumps.
This results in muscle
relaxation.

Mitochondrion

Transverse
tubules
Sarcoplasmic
reticulum

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Neuromuscular junction

Junction of motor neurons axon and muscle fiber

Axon divides into terminals containing vesicles of


acetylcholine

Region of muscle fiber under axon terminal is folded into


junctional folds to increase surface area

ACh receptor is ligand-gated ion channel

Na+ flows into muscle cell leading to depolarization


and an action potential
30

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Myelin
Skeletal
muscle
fibers
Motor
neuron
Neuromuscular
junction

Axon terminal

(a) SEM of neuromuscular junction


Plasma
membrane of
muscle fiber
Axon terminal
Synaptic
cleft

Junctional
Junctional
folds
folds with
with ACh
ACh
receptors
receptors

Synaptic
vesicles
containing ACh
Na+
ACh receptor

Motor
end plate

ACh

Acetylcholinesterase

(b) Structures of, and events at, the neuromuscular junction


(only part of the motor neuron is shown)
Dr. Donald Fawcett/Photo Researchers, Inc.

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Skeletal Muscle Function

Different muscle fibers contain forms of myosin that differ


in the maximal rates at which they can hydrolyze ATP

Fast fibers contain myosin with high ATPase activity

Slow fibers have myosin with a lower ATPase activity

Maximal force produced by each is the same,


only speed varies

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Oxidative fibers
Contain

numerous mitochondria and have a high


capacity for oxidative phosphorylation

Depends

on blood flow to deliver oxygen and


nutrients for ATP production

Contain

large amounts of myoglobin as an


intracellular reservoir of oxygen

Glycolytic fibers
Few

mitochondria but a high concentration of


glycolytic enzymes and large stores of glycogen

Contain

little myoglobin makes them pale or white


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Three major types


1.

2.

3.

Slow-oxidative fibers

Low rates of myosin ATP hydrolysis but makes


large amounts of ATP

Used for prolonged, regular activity

Fast-oxidative fibers

High myosin activity, makes large amounts of ATP

Suited for rapid actions

Fast-glycolytic fibers

High myosin activity but cannot make as much ATP

Suited for rapid, intense actions but fatigues quickly

Exercise

Increased amounts of exercise can produce an


increase in the size of muscle fibers and their
capacity for ATP production
Increase

in muscle size due to increase in size of


individual fibers

Atrophy is the reduction in size of a muscle


Happens

if neurons at neuromuscular junction


become nonfunctional
36

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32 m

32 m

37

Factors That Influence Strength of Muscle


Contraction

Contraction of a whole muscle depends on the


types of muscle fibers making up the muscle, the
number of muscle fibers contracting, the tension
developed by each fiber, frequency of stimulation,
and the condition of the muscle
Aerobic training increases capillary density, number
of mitochondria, and myoglobin content of muscle
fibers
Strength training increases the number of filaments
in muscle fibers, which increases the size of the
muscle

Factors That Influence Strength of Muscle


Contraction (cont.)

The more muscle fibers that contract, the greater


the tension in the muscle in a motor unit, a
motor neuron is functionally connected with
several to hundreds of muscle fibers

Each junction of a motor neuron with a muscle


fiber is called a neuromuscular junction

Messages from the brain or spinal cord activate


motor units the more motor units recruited, the
stronger the contractions

Factors That Influence Strength of Muscle


Contraction (cont.)

In response to a single, brief electrical stimulus,


skeletal muscle contracts with a single, quick simple
twitch
When a series of separate stimuli arrive very closely
together, the twitches add together (summation),
resulting in a smooth, sustained contraction (tetanus)
When you are not moving, your muscles are in a state
of partial contraction (muscle tone), an unconscious
process that keeps muscles prepared for action

Contracting muscle
exerts a force on bones
through its connecting
tendons
Contracting muscle
exerts only a pulling force
Muscles that bend a limb
are flexors
Muscles that straighten a
limb are extensors
Groups of muscles with
oppositely directed
motions at a joint are
antagonists

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Extensors relaxed
(quadriceps)

Extensors
contracted

Flexors
contracted
(hamstring)

Flexors relaxed

42

Animal locomotion

Water
Greatest

challenge is the density of water


Resistance increases exponentially as speed
increases
Streamlined bodies reduce drag and make swimming
more efficient
Energetic advantage is that swimmers do not need to
provide lift to overcome gravity
Swimming similar among many vertebrates
Invertebrates use means other than swimming
squid propulsion, move passively on current, crawl on
rocks

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Locomotion on land walking and running


Most

energetically costly means of locomotion

Even flying is not as costly

Gravity

must be overcome at each step

Accelerating

and decelerating with each step is more

important
Most

animals limit ground contact to reduce friction

Except mollusks on mucus and snakes undulating

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Locomotion in air flying


Evolved

on FOUR occasions

Pterosaurs, insects, birds, and mammals (bats)

advantages escape, scan large areas,


inhabit inaccessible areas

Numerous

Mechanics

require overcoming gravity and


air resistance

Resistance

reduced by streamlined bodies

Lift

and thrust in vertebrates provided by pectoral


and back muscles
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Bird and bat wings are modifications of the


forelimbs
Bat wings more maneuverable with fingers
at the end of forelimb/wings but cant glide
for long
Birds have less control but large birds can
glide for long periods of time

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Impact on public health

Rickets
Improper

mineral deposition
Usually due to inadequate
dietary calcium intake or
inadequate absorption of
calcium from the small
intestine
Prevented or treated with
vitamin D
48

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Osteoporosis
Both

mineral and organic


portions of bone reduced

Occurs

when normal
balance between bone
formation and breakdown
is disrupted

Can

be minimized with
adequate calcium intake,
weight-bearing exercise,
and adequate dietary
Vitamin D

2 m
32 m

32 m
2 m

(b) Histologic appearance of


normal bone (top) and
bone from a person with
osteoporosis (bottom)
b: Tim Arnett, University College London

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Myasthenia gravis MS
Characterized

by skeletal muscle fatigue and

weakness
Affects

10,000 30,000 Americans

Bodys

immune system produces antibodies that bind


to and inactivate ACh receptors on skeletal muscles
an autoimmune disease

Treatments

range from enzyme inhibitors that allow


ACh persist, to immune system suppression, to
plasmapheresis to remove antibodies from the blood

50

Muscular dystrophy
Group

of diseases affecting 1 in 3500 American


males (less common in females)

Progressive

degeneration of skeletal and cardiac


muscles, ultimately leading to death

Most

common form is Duchenne muscular


dystrophy sex-linked recessive disorder

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