CH 10 Lecture Presentation

Chapter 10
Muscle Tissue
Lecture Presentation by
Lee Ann Frederick
University of Texas at Arlington
© 2015 Pearson Education, Inc.

10-1 An Introduction to Muscle Tissue
• Learning Outcomes
• 10-1 Specify the functions of skeletal muscle
tissue.
• 10-2 Describe the organization of muscle at the
tissue level.
• 10-3 Describe the characteristics of skeletal
muscle fibers, and identify the structural
components of a sarcomere.
• 10-4 Identify the components of the
neuromuscular junction, and summarize the
events involved in the neural control of
skeletal muscle contraction and relaxation.
• 10-5 Describe the mechanism responsible for
tension production in a muscle fiber, and
compare the different types of muscle
contraction.
• 10-6 Describe the mechanisms by which muscle
fibers obtain the energy to power
contractions.
• 10-7 Relate the types of muscle fibers to muscle
performance, and distinguish between
aerobic and anaerobic endurance.

• 10-8 Identify the structural and functional
differences between skeletal muscle fibers
and cardiac muscle cells.
• 10-9 Identify the structural and functional
differences between skeletal muscle fibers
and smooth muscle cells, and discuss the
roles of smooth muscle tissue in systems
throughout the body.

An Introduction to Muscle Tissue
• Muscle Tissue
• A primary tissue type, divided into:
• Skeletal muscle tissue
• Cardiac muscle tissue
• Smooth muscle tissue

10-1 Functions of Skeletal Muscle Tissue
• Skeletal Muscles
• Are attached to the skeletal system
• Allow us to move
• The muscular system
• Includes only skeletal muscles

10-1 Functions of Skeletal Muscle Tissue
• Six Functions of Skeletal Muscle Tissue

1. Produce skeletal movement
2. Maintain posture and body position
3. Support soft tissues
4. Guard entrances and exits
5. Maintain body temperature
6. Store nutrient reserves

10-2 Organization of Muscle
• Skeletal Muscle
• Muscle tissue (muscle cells or fibers)
• Connective tissues
• Nerves
• Blood vessels

• Organization of Connective Tissues

• Muscles have three layers of connective tissues
1. Epimysium
2. Perimysium
3. Endomysium

• Epimysium
• Exterior collagen layer
• Connected to deep fascia
• Separates muscle from surrounding tissues

• Perimysium
• Surrounds muscle fiber bundles (fascicles)
• Contains blood vessel and nerve supply to
fascicles

• Endomysium
• Surrounds individual muscle cells (muscle fibers)
• Contains capillaries and nerve fibers contacting
muscle cells
• Contains myosatellite cells (stem cells) that repair
damage

Figure 10-1 The Organization of Skeletal Muscles (Part 1 of 3).
Skeletal Muscle (organ)
Epimysium Perimysium Endomysium Nerve
Muscle Muscle Blood

fascicle fibers vessels
Epimysium
Blood vessels
and nerves
Tendon
Endomysium
Perimysium

Muscle Fascicle (bundle of fibers)
Perimysium
Muscle fiber
Epimysium
Blood vessels
and nerves
Endomysium
Tendon
Endomysium
Perimysium

Muscle Fiber (cell)
Capillary Myofibril Endomysium
Sarcoplasm
Epimysium
Blood vessels Mitochondrion

and nerves
Myosatellite
cell
Sarcolemma
Nucleus
Tendon
Axon of neuron
Endomysium
Perimysium

• Organization of Connective Tissues

• Muscle Attachments
• Endomysium, perimysium, and epimysium come
together:
• At ends of muscles
• To form connective tissue attachment to bone matrix
• i.e., tendon (bundle) or aponeurosis (sheet)

• Blood Vessels and Nerves

• Muscles have extensive vascular systems that:
• Supply large amounts of oxygen
• Supply nutrients
• Carry away wastes
• Skeletal muscles are voluntary muscles, controlled
by nerves of the central nervous system (brain
and spinal cord)

10-3 Characteristics of Skeletal Muscle
Fibers
• Skeletal Muscle Cells
• Are very long
• Develop through fusion of mesodermal cells
(myoblasts)
• Become very large
• Contain hundreds of nuclei

Figure 10-2 The Formation of a Multinucleate Skeletal Muscle Fiber.
Muscle fibers develop through

the fusion of embryonic
cells called myoblasts.
Myoblasts
a A muscle
Muscle fiber LM × 612
fiber forms by
the fusion of
myoblasts. Sarcolemma Striations Nuclei
Myofibrils
Myosatellite cell
Nuclei
Mitochondria
Immature
muscle fiber
b A diagrammatic view and a

Myosatellite cell micrograph of one muscle
fiber.
Up to 30 cm
in length
Mature muscle fiber

Figure 10-2a The Formation of a Multinucleate Skeletal Muscle Fiber.
Muscle fibers develop through

the fusion of embryonic
cells called myoblasts.
Myoblasts
a A muscle
fiber forms by
the fusion of
myoblasts.
Myosatellite cell
Nuclei
Immature
muscle fiber
Myosatellite cell
Up to 30 cm
in length
Mature muscle fiber

Figure 10-2b The Formation of a Multinucleate Skeletal Muscle Fiber.
Muscle fiber LM × 612
Sarcolemma Striations Nuclei
Myofibrils
Mitochondria
b A diagrammatic view and a

micrograph of one muscle
fiber.
Fibers
• The Sarcolemma and Transverse Tubules
• The sarcolemma
• The cell membrane of a muscle fiber (cell)
• Surrounds the sarcoplasm (cytoplasm of muscle
fiber)
• A change in transmembrane potential begins
contractions

Fibers
• The Sarcolemma and Transverse Tubules
• Transverse tubules (T tubules)
• Transmit action potential through cell
• Allow entire muscle fiber to contract simultaneously
• Have same properties as sarcolemma

Fibers
• Myofibrils
• Lengthwise subdivisions within muscle fiber
• Made up of bundles of protein filaments
(myofilaments)
• Myofilaments are responsible for muscle
contraction
• Types of myofilaments:
• Thin filaments
• Made of the protein actin
• Thick filaments
• Made of the protein myosin

Fibers
• The Sarcoplasmic Reticulum (SR)
• A membranous structure surrounding each
myofibril
• Helps transmit action potential to myofibril
• Similar in structure to smooth endoplasmic
reticulum
• Forms chambers (terminal cisternae) attached to
T tubules

Fibers
• The Sarcoplasmic Reticulum (SR)
• Triad
• Is formed by one T tubule and two terminal
cisternae
• Cisternae
• Concentrate Ca2+ (via ion pumps)
• Release Ca2+ into sarcomeres to begin muscle
contraction

Figure 10-3 The Structure and Internal Organization of a Skeletal Muscle Fiber.
Myofibril
Muscle fiber
Sarcolemma Nuclei
Sarcoplasm
Mitochondria
Terminal cisterna
Sarcolemma
Sarcolemma
Sarcoplasm
Myofibril
Myofibrils
Thin filament
Thick filament
Triad Sarcoplasmic T tubules
reticulum

Figure 10-3 The Structure and Internal Organization of a Skeletal Muscle Fiber (Part 1 of 4).
Myofibril
Muscle fiber
Sarcolemma Nuclei
Sarcoplasm

Mitochondria
Terminal cisterna
Sarcolemma
Sarcolemma
Sarcoplasm
Myofibril
Myofibrils
Thin filament
Thick filament
reticulum

Mitochondria
Sarcolemma
Myofibril
Thin filament
Thick filament

Terminal cisterna
Sarcolemma
Sarcoplasm
Myofibrils

reticulum

10-3 Structural Components of a Sarcomere
• Sarcomeres
• The contractile units of muscle
• Structural units of myofibrils
• Form visible patterns within myofibrils
• A striped or striated pattern within myofibrils
• Alternating dark, thick filaments (A bands) and
light, thin filaments (I bands)

• Sarcomeres
• The A Band
• M line
• The center of the A band
• At midline of sarcomere
• The H Band
• The area around the M line
• Has thick filaments but no thin filaments
• Zone of overlap
• The densest, darkest area on a light micrograph
• Where thick and thin filaments overlap

• Sarcomeres
• The I Band
• Z lines
• The centers of the I bands
• At two ends of sarcomere
• Titin
• Are strands of protein
• Reach from tips of thick filaments to the Z line
• Stabilize the filaments

Figure 10-4a Sarcomere Structure, Longitudinal Views.
I band A band
H band Z line Titin
a A longitudinal view of a
sarcomere, showing bands
of thick and thin filaments
Zone of overlap M line Thin Thick

filament filament
Sarcomere

Figure 10-4b Sarcomere Structure, Longitudinal Views.
I band A band
H band Z line
b A corresponding
longitudinal section of a
sarcomere in a myofibril
from a muscle fiber in
the gastrocnemius (calf) Myofibril TEM × 64,000
muscle of the leg
Z line Zone of overlap M line
Sarcomere

Figure 10-5 Sarcomere Structure, Superficial and Cross-Sectional Views.
Sarcomere
Myofibril
a A superficial view
of a sarcomere
Thin Thick
filament filament
Actinin Thin filaments Titin Thick filaments

filaments filament
Attachment
of titin
Z line I band M line H band Zone of overlap

b Cross-sectional views of different
regions of a sarcomere

Figure 10-6 Levels of Functional Organization in a Skeletal Muscle.
Skeletal Muscle Myofibril
Surrounded by: Surrounded by:

Epimysium Sarcoplasmic
Epimysium reticulum
Contains:
Muscle fascicles Consists of:
Sarcomeres
(Z line to Z line)
Sarcomere
I band A band
Muscle Fascicle
Contains:
Thick filaments
Surrounded by:
Perimysium Thin filaments
Perimysium
Contains:
Muscle fibers
Z line M line Titin Z line
H band
Muscle Fiber
Surrounded by:
Endomysium
Endomysium
Contains:
Myofibrils

Figure 10-6 Levels of Functional Organization in a Skeletal Muscle (Part 1 of 5).
Skeletal Muscle
Surrounded by:
Epimysium
Epimysium
Contains:
Muscle fascicles

Muscle Fascicle
Surrounded by:
Perimysium
Perimysium
Contains:
Muscle fibers

Muscle Fiber
Surrounded by:
Endomysium
Endomysium
Contains:
Myofibrils

Myofibril
Surrounded by:
Sarcoplasmic
reticulum
Consists of:
Sarcomeres
(Z line to Z line)

Sarcomere
I band A band
Contains:
Thick filaments
Thin filaments
Z line M line Titin Z line

H band

• Thin Filaments
• F-actin (filamentous actin)
• Is two twisted rows of globular G-actin
• The active sites on G-actin strands bind to myosin
• Nebulin
• Holds F-actin strands together

• Thin Filaments
• Tropomyosin
• Is a double strand
• Prevents actin–myosin interaction
• Troponin
• A globular protein
• Binds tropomyosin to G-actin
• Controlled by Ca2+

Figure 10-7ab Thin and Thick Filaments.
Sarcomere
H band Actinin Z line Titin
Myofibril a The gross structure of a thin

filament, showing the attachment
at the Z line
Troponin Active site Nebulin Tropomyosin G-actin

Z line M line molecules
F-actin
strand
b The organization of G-actin subunits in an F-actin strand,

and the position of the troponin–tropomyosin complex

• Initiating Contraction
• Ca2+ binds to receptor on troponin molecule
• Troponin–tropomyosin complex changes
• Exposes active site of F-actin

• Thick Filaments
• Contain about 300 twisted myosin subunits
• Contain titin strands that recoil after stretching
• The mysosin molecule
• Tail
• Binds to other myosin molecules
• Head
• Made of two globular protein subunits
• Reaches the nearest thin filament

Figure 10-7cd Thin and Thick Filaments.
Titin
c The structure of thick

filaments, showing M line Myosin
Myosin tail
the orientation of the head
myosin molecules
Hinge
d The structure of a myosin molecule

• Myosin Action
• During contraction, myosin heads:
• Interact with actin filaments, forming cross-bridges
• Pivot, producing motion

• Sliding Filaments and Muscle Contraction

• Sliding filament theory
• Thin filaments of sarcomere slide toward M line,
alongside thick filaments
• The width of A zone stays the same
• Z lines move closer together

Figure 10-8a Changes in the Appearance of a Sarcomere during the Contraction of a Skeletal Muscle Fiber.
Myofibril at rest
I band A band
Z line H band Z line

a A relaxed sarcomere showing location of
the A band, Z lines, and I band.

Figure 10-8b Changes in the Appearance of a Sarcomere during the Contraction of a Skeletal Muscle Fiber.
Contracted myofibril
I band A band
Z line H band Z line

b During a contraction, the A band stays the same width,
but the Z lines move closer together and the I band
gets smaller. When the ends of a myofibril are free to
move, the sarcomeres shorten simultaneously and the
ends of the myofibril are pulled toward its center.
• Skeletal Muscle Contraction

• The process of contraction
• Neural stimulation of sarcolemma
• Causes excitation–contraction coupling
• Muscle fiber contraction
• Interaction of thick and thin filaments
• Tension production

10-4 Components of the Neuromuscular
Junction
• The Control of Skeletal Muscle Activity
• The neuromuscular junction (NMJ)
• Special intercellular connection between the
nervous system and skeletal muscle fiber
• Controls calcium ion release into the sarcoplasm

A&P FLIX Events at the Neuromuscular
Junction

Figure 10-9 Events at the Neuromuscular Junction (Part 3 of 9).
A single axon may branch to control more than one

Motor neuron
skeletal muscle fiber, but each muscle fiber has only
one neuromuscular junction (NMJ). At the NMJ, the Path of electrical impulse
axon terminal of the neuron lies near the motor end (action potential)
plate of the muscle fiber. Axon
Neuromuscular
junction
Axon
terminal
SEE BELOW
Sarcoplasmic Motor
reticulum end plate
Myofibril
Motor end plate

1
The cytoplasm of the axon
terminal contains vesicles filled
with molecules of acetylcholine,
or ACh. Acetylcholine is a
neurotransmitter, a chemical
released by a neuron to change
the permeability or other
properties of another cell’s plasma
membrane. The synaptic cleft and
the motor end plate contain
molecules of the enzyme
acetylcholinesterase (AChE),
which breaks down ACh.
Vesicles ACh
The synaptic cleft is a

narrow space that separates
the axon terminal of the
neuron from the opposing
motor end plate.
Junctional AChE
fold of
motor end plate
2
The stimulus for ACh release
is the arrival of an electrical
impulse, or action potential,
at the axon terminal. An action
potential is a sudden change in
the membrane potential that
travels along the length of the
axon.
Arriving action
potential

3
When the action potential
reaches the neuron’s axon
terminal, permeability
changes in its membrane
trigger the exocytosis of ACh
into the synaptic cleft.
Exocytosis occurs as vesicles
fuse with the neuron’s plasma
membrane.
Motor
end plate

4
ACh molecules diffuse across the
synaptic cleft and blind to ACh
receptors on the surface of the
motor end plate. ACh binding
alters the membrane’s permeabil-
ity to sodium ions. Because the
extracellular fluid contains a high
concentration of sodium ions,
and sodium ion concentration
inside the cell is very low, sodium
ions rush into the cytosol.
Na+
Na+
Na+
ACh
receptor site
5
The sudden inrush of sodium ions
results in the generation of an
action potential in the sarcolemma.
ACh is removed from the
synaptic cleft in two ways. ACh
either diffuses away from the
synapse, or it is broken down by
AChE into acetic acid and choline.
This removal inactivates the ACh
receptor sites. The muscle fiber
pictured above indicates the
propagation of the action
potential along the sarcolemma.
Action
potential
Break down AChE

of ACh

10-4 Components of the Neuromuscular
Junction
• Excitation–Contraction Coupling
• Action potential reaches a triad
• Releasing Ca2+
• Triggering contraction
• Requires myosin heads to be in “cocked” position
• Loaded by ATP energy

A&P FLIX Excitation-Contraction Coupling

Figure 10-10 Excitation-Contraction Coupling.




10-4 Skeletal Muscle Contraction
• The Contraction Cycle

1. Contraction Cycle Begins
2. Active-Site Exposure
3. Cross-Bridge Formation
4. Myosin Head Pivoting
5. Cross-Bridge Detachment
6. Myosin Reactivation

A&P FLIX The Cross Bridge Cycle

Figure 10-11 The Contraction Cycle and Cross-Bridge Formation (Part 3 of 10).
1
Contraction Cycle Begins
The contraction cycle involves a
series of interrelated steps. It begins
with the arrival of calcium ions
(Ca2+) within the zone of overlap in
a sarcomere.
ADP
+ Ca2+ Myosin head
P
Troponin
Tropomyosin Actin
ADP Ca2+
P+

2
Active-Site Exposure
Calcium ions bind to troponin,
weakening the bond between
actin and the troponin–tropomyosin
complex. The troponin molecule
then changes position, rolling the
tropomyosin molecule away from
the active sites on actin and
allowing interaction with the
energized myosin heads.
ADP
+ P
Cytosol
Ca2+
Ca2+
Active
site ADP
P+

3
Cross-Bridge Formation
Once the active sites are
exposed, the energized myosin
heads bind to them, forming
cross-bridges.
ADP
+ Ca2+
P
Ca2+
ADP
P+

4
Myosin Head Pivoting
After cross-bridge formation,
the energy that was stored in the
resting state is released as the
myosin head pivots toward the M
line. This action is called the power
stroke; when it occurs,
the bound ADP and phosphate
group are released.
ADP + P
Ca2+
Ca2+
ADP + P

5
Cross-Bridge Detachment
When another ATP binds to the
myosin head, the link between the
myosin head and the active site on
the actin molecule is broken. The
active site is now exposed and able
to form another cross-bridge.
ATP
Ca2+
Ca2+
ATP

6
Myosin Reactivation
Myosin reactivation occurs
when the free myosin head
splits ATP into ADP and P.
The energy released is used
to recock the myosin head.
ADP
+ P
Ca2+
Ca2+
ADP
P+

In the resting sarcomere,

each myosin head is already
“energized”—charged with
the energy that will be used
to power a contraction. Each
myosin head points away
from the M line. In this
position, the myosin head is
“cocked” like the spring in a
mousetrap. Cocking the myosin M line Zone of Overlap
head requires energy, which is (shown in
obtained by breaking down sequence above)
ATP; in doing so, the myosin
head functions as ATPase, an
enzyme that breaks down ATP.
At the start of the contraction
cycle, the breakdown products,
ADP and phosphate (represent-
ed as P), remain bound to the
myosin head.

The entire cycle is repeated several

times each second, as long as Ca2+
concentrations remain elevated and
ATP reserves are sufficient. Calcium
ion levels will remain elevated only
as long as action potentials continue
to pass along the T tubules and stimulate
the terminal cisternae. Once that stimulus
is removed, the calcium channels in the
SR close and calcium ion pumps pull
Ca2+ from the cytosol and store it
within the terminal cisternae. Troponin
molecules then shift position, swinging
the tropomyosin strands over the active
sites and preventing further cross-bridge
formation.

10-4 Skeletal Muscle Contraction
• Fiber Shortening
• As sarcomeres shorten, muscle pulls together,
producing tension
• Muscle shortening can occur at both ends of the
muscle, or at only one end of the muscle
• This depends on the way the muscle is attached at
the ends

Figure 10-12 Shortening during a Contraction.
a When both ends are free to move, the ends of a contracting

muscle fiber move toward the center of the muscle fiber.
b When only one end of a myofibril is fixed in position,

the free end is pulled toward the fixed end.

10-4 Skeletal Muscle Relaxation
• Relaxation
• Contraction duration
• Depends on:
• Duration of neural stimulus
• Number of free calcium ions in sarcoplasm
• Availability of ATP

10-4 Skeletal Muscle Relaxation
• Relaxation
• Ca2+ concentrations fall
• Ca2+ detaches from troponin
• Active sites are re-covered by tropomyosin
• Rigor Mortis
• A fixed muscular contraction after death
• Caused when:
• Ion pumps cease to function; ran out of ATP
• Calcium builds up in the sarcoplasm

10-4 Skeletal Muscle Contraction and
Relaxation
• Summary
• Skeletal muscle fibers shorten as thin filaments
slide between thick filaments
• Free Ca2+ in the sarcoplasm triggers contraction
• SR releases Ca2+ when a motor neuron stimulates
the muscle fiber
• Contraction is an active process
• Relaxation and return to resting length are passive

Figure 10-13 Steps Involved in Skeletal Muscle Contraction and Relaxation (Part 1 of 2).
Steps That Initiate a Muscle Contraction
1 ACh released Axon

Sarcolemma
terminal
ACh is released at the neuromuscular junction and binds to
ACh receptors on the sarcolemma.
Cytosol
2 Action potential reaches T tubule T tubule
An action potential is generated and spreads across the

membrane surface of the muscle fiber and along the T Sarcoplasmic reticulum
tubules.
3 Sarcoplasmic reticulum releases Ca2+ Ca2+

The sarcoplasmic reticulum releases stored calcium ions.
Actin
4 Active site exposure and cross-bridge formation

Myosin
Calcium ions bind to troponin, exposing the active sites

on the thin filaments. Cross-bridges form when myosin
heads bind to those active sites.
5 Contraction cycle begins

The contraction cycle begins as repeated cycles of
cross-bridge binding, pivoting, and detachment occur—all
powered by ATP.

Figure 10-13 Steps Involved in Skeletal Muscle Contraction and Relaxation (Part 2 of 2).
Steps That End a Muscle Contraction
Axon
6 ACh is broken down terminal Sarcolemma
ACh is broken down by acetylcholinesterase (AChE),

ending action potential generation
Cytosol
T tubule
7 Sarcoplasmic reticulum reabsorbs Ca2+
As the calcium ions are reabsorbed, their concentration in Sarcoplasmic reticulum
the cytosol decreases.
8 Active sites covered, and cross-bridge formation ends

Ca2+
Without calcium ions, the tropomyosin returns to its normal Actin

position and the active sites are covered again.
Myosin
9 Contraction ends
Without cross-bridge formation, contraction ends.
10 Muscle relaxation occurs

The muscle returns passively to its resting length.

10-5 Tension Production and Contraction
Types
• Tension Production by Muscles Fibers
• As a whole, a muscle fiber is either contracted or
relaxed
• Depends on:
• The number of pivoting cross-bridges
• The fiber’s resting length at the time of stimulation
• The frequency of stimulation

Types
• Tension Production by Muscles Fibers
• Length–Tension Relationships
• Number of pivoting cross-bridges depends on:
• Amount of overlap between thick and thin fibers
• Optimum overlap produces greatest amount of
tension
• Too much or too little reduces efficiency
• Normal resting sarcomere length
• Is 75 to 130 percent of optimal length

Figure 10-14 The Effect of Sarcomere Length on Active Tension.
Maximum tension is produced

when the zone of overlap is large
but the thin filaments do not extend
At short resting lengths, thin across the sarcomere’s center.
filaments extending across the
center of the sarcomere interfere If the sarcomeres are stretched too far, the
with the normal orientation of thick zone of overlap is reduced or disappears,
and thin filaments, decreasing and cross-bridge interactions are reduced
tension production. or cannot occur.
100
Tension (percent of maximum)
80
60
When the thick When the zone of overlap is reduced to

filaments contact the 40 zero, thin and thick filaments cannot
Z lines, the sarcomere interact at all. The muscle fiber cannot
cannot shorten—the produce any active tension, and a
20 Normal
myosin heads cannot contraction cannot occur. Such extreme
range
pivot and tension stretching of a muscle fiber is normally
cannot be produced. prevented by titin filaments (which tie the
0
thick filaments to the Z lines) and by the
1.2 μm 1.6 μm 2.6 μm 3.6 μm surrounding connective tissues.
Sarcomere length Sarcomere length
decreases increases
Optimal resting length:
The normal range of sarcomere
lengths in the body is 75 to 130
percent of the optimal length.

Types
• Tension Production by Muscle Fibers
• The Frequency of Stimulation
• A single neural stimulation produces:
• A single contraction or twitch
• Which lasts about 7–100 msec.
• Sustained muscular contractions
• Require many repeated stimuli

Types
• Twitches
1. Latent period
• The action potential moves through sarcolemma
• Causing Ca2+ release
2. Contraction phase
• Calcium ions bind
• Tension builds to peak
3. Relaxation phase
• Ca2+ levels fall
• Active sites are covered and tension falls to resting
levels

Figure 10-15a The Development of Tension in a Twitch.
Eye muscle
Gastrocnemius
Soleus
Tension
0 10 20 30 40 50 60 70 80 90 100
Time (msec)
Stimulus
a A myogram showing differences in tension over
time for a twitch in different skeletal muscles.
Figure 10-15b The Development of Tension in a Twitch.
Maximum tension
development
Tension
Stimulus
Time (msec) 0 5 10 20 30 40
Resting Latent Contraction Relaxation

phase period phase phase
b The details of tension over time for a single twitch
in the gastrocnemius muscle. Notice the presence
of a latent period, which corresponds to the time
needed for the conduction of an action potential
and the subsequent release of calcium ions by the
sarcoplasmic reticulum.
Types
• Treppe
• A stair-step increase in twitch tension
• Repeated stimulations immediately after relaxation
phase
• Stimulus frequency 50/second
• Causes a series of contractions with increasing
tension

Types
• Wave summation
• Increasing tension or summation of twitches
• Repeated stimulations before the end of relaxation
phase
• Stimulus frequency 50/second
• Causes increasing tension or summation of
twitches

Figure 10-16ab Effects of Repeated Stimulations.
= Stimulus
Maximum tension (in tetanus)
Tension
Maximum tension (in treppe)
Time Time
a Treppe. Treppe is an increase in b Wave summation. Wave summation
peak tension with each successive occurs when successive stimuli
stimulus delivered shortly after the arrive before the relaxation phase
completion of the relaxation phase of has been completed.
the preceding twitch.

Types
• Incomplete tetanus
• Twitches reach maximum tension
• If rapid stimulation continues and muscle is not
allowed to relax, twitches reach maximum level of
tension
• Complete tetanus
• If stimulation frequency is high enough, muscle
never begins to relax, and is in continuous
contraction

Figure 10-16cd Effects of Repeated Stimulations.
Maximum tension (in tetanus)

Tension
Time Time
c Incomplete tetanus. Incomplete d Complete tetanus. During
tetanus occurs if the stimulus complete tetanus, the stimulus
frequency increases further. Tension frequency is so high that the
production rises to a peak, and the relaxation phase is eliminated.
periods of relaxation are very brief. Tension plateaus at maximum levels.

Types
• Tension Production by Skeletal Muscles
• Depends on:
• Internal tension produced by muscle fibers
• External tension exerted by muscle fibers on elastic
extracellular fibers
• Total number of muscle fibers stimulated

Types
• Motor Units and Tension Production
• Motor units in a skeletal muscle:
• Contain hundreds of muscle fibers
• That contract at the same time
• Controlled by a single motor neuron

Types
• Recruitment (multiple motor unit summation)
• In a whole muscle or group of muscles, smooth
motion and increasing tension are produced by
slowly increasing the size or number of motor
units stimulated
• Maximum tension
• Achieved when all motor units reach tetanus
• Can be sustained only a very short time

Figure 10-17a The Arrangement and Activity of Motor Units in a Skeletal Muscle.
Axons of
motor neurons
SPINAL CORD
Motor
nerve
KEY
Muscle fibers
Motor unit 1
Motor unit 2
Motor unit 3
a Muscle fibers of different motor units are

intermingled, so the forces applied to the
tendon remain roughly balanced regardless
of which motor units are stimulated.
Figure 10-17b The Arrangement and Activity of Motor Units in a Skeletal Muscle.
Tension in tendon
Motor Motor Motor

unit 1 unit 2 unit 3
Tension
Time
b The tension applied to the tendon
remains relatively constant, even
though individual motor units cycle
between contraction and relaxation.

Types
• Sustained tension
• Less than maximum tension
• Allows motor units to rest in rotation
• Muscle tone
• The normal tension and firmness of a muscle at
rest
• Muscle units actively maintain body position,
without motion
• Increasing muscle tone increases metabolic energy
used, even at rest

Types
• Contractions are classified based on pattern of
tension production
• Isotonic contraction
• Isometric contraction

Types
• Isotonic Contraction
• Skeletal muscle changes length
• Resulting in motion
• If muscle tension  load (resistance):
• Muscle shortens (concentric contraction)
• If muscle tension  load (resistance):
• Muscle lengthens (eccentric contraction)

Figure 10-18a Concentric, Eccentric, and Isometric Contractions.
Tendon Amount of Muscle

Muscle 4 load relaxes
tension Peak tension
(kg) 2 production
Muscle
contracts 0
(concentric Contraction
contraction) begins Resting length
100
Muscle
2 kg 90 length
(percent
2 kg 80 of resting
length)
70
a
Time
In this experiment, a muscle is attached to a weight one-half its peak
tension potential. On stimulation, it develops enough tension to lift the
weight. Tension remains constant for the duration of the contraction,
although the length of the muscle changes. This is an example of
isotonic contraction.

Figure 10-18b Concentric, Eccentric, and Isometric Contractions.
When the eccentric contraction ends, the

unopposed load stretches the muscle until
either the muscle tears, a tendon breaks, or
the elastic recoil of the skeletal muscle is
sufficient to oppose the load.
Support removed 4 140

when contraction Muscle
begins Peak tension 130
tension 2
(eccentric contraction) (kg) production
120 Muscle
0 length
110 (percent
Support removed,
of resting
contraction begins 100 length)
6 kg Resting length
90
80
6 kg
70
b In this eccentric contraction, the muscle elongates as it generates tension. Time

Types
• Isometric Contraction
• Skeletal muscle develops tension, but is
prevented from changing length
• iso-  same, metric  measure

Figure 10-18c Concentric, Eccentric, and Isometric Contractions.
Amount of load
6
Muscle
Muscle relaxes
4
tension
(kg) Peak tension
Muscle 2
contracts production
(isometric
0
contraction)
Contraction
begins Length unchanged
100
Muscle
90 length
6 kg 6 kg (percent
80 of resting
length)
70
c The same muscle is attached to a weight that exceeds its peak Time
tension capabilities. On stimulation, tension will rise to a peak,
but the muscle as a whole cannot shorten. This is an isometric
contraction.

Types
• Load and Speed of Contraction
• Are inversely related
• The heavier the load (resistance) on a muscle:
• The longer it takes for shortening to begin
• And the less the muscle will shorten

Figure 10-19 Load and Speed of Contraction.
Distance shortened
Small load
Intermediate load
Large load
0 20 40 60 80 100 120 140

Time (msec)
Stimulus
Types
• Muscle Relaxation and the Return to Resting
Length
• Elastic forces
• The pull of elastic elements (tendons and
ligaments)
• Expands the sarcomeres to resting length
• Opposing muscle contractions
• Reverse the direction of the original motion
• Are the work of opposing skeletal muscle pairs

Types
• Muscle Relaxation and the Return to Resting
Length
• Gravity
• Can take the place of opposing muscle contraction
to return a muscle to its resting state

10-6 Energy to Power Contractions
• ATP Provides Energy for Muscle Contraction

• Sustained muscle contraction uses a lot of ATP
energy
• Muscles store enough energy to start contraction
• Muscle fibers must manufacture more ATP as
needed

• ATP and CP Reserves

• Adenosine triphosphate (ATP)
• The active energy molecule
• Creatine phosphate (CP)
• The storage molecule for excess ATP energy in
resting muscle
• Energy recharges ADP to ATP
• Using the enzyme creatine kinase (CK)
• When CP is used up, other mechanisms generate
ATP

• ATP Generation
• Cells produce ATP in two ways
1. Aerobic metabolism of fatty acids in the
mitochondria
2. Anaerobic glycolysis in the cytoplasm

• Aerobic Metabolism
• Is the primary energy source of resting muscles
• Breaks down fatty acids
• Produces 34 ATP molecules per glucose molecule
• Glycolysis
• Is the primary energy source for peak muscular
activity
• Produces two ATP molecules per molecule of
glucose
• Breaks down glucose from glycogen stored in
skeletal muscles
Table 10-1 Sources of Energy in a Typical Muscle Fiber.

• Energy Use and the Level of Muscular Activity

• Skeletal muscles at rest metabolize fatty acids and
store glycogen
• During light activity, muscles generate ATP
through anaerobic breakdown of carbohydrates,
lipids, or amino acids
• At peak activity, energy is provided by anaerobic
reactions that generate lactic acid as a by-
product

Figure 10-20 Muscle Metabolism (Part 1 of 3).
Muscle Metabolism in a Resting Muscle Fiber
• In a resting skeletal muscle, the demand for ATP is low, Fatty acids O2 G Blood vessels
and there is more than enough oxygen available for
mitochondria to meet that demand.
• Resting muscle fibers absorb fatty acids, which are broken

down in the mitochondria creating a surplus of ATP. Glucose Glycogen
• Some mitochondrial ATP is used to convert absorbed
glucose to glycogen. ADP ADP
CP
• Mitochondrial ATP is also used to convert creatine to Mitochondria ATP
creatine phosphate (CP).
• This results in the buildup of energy reserves (glycogen CO2 Creatine
and CP) in the muscle.

Muscle Metabolism during Moderate Activity
• During moderate levels of activity, the demand for ATP Fatty acids
O2
increases.
• There is still enough oxygen for the mitochondria to meet
that demand, but no excess ATP is produced.
Glucose Glycogen
• The muscle fiber now relies primarily on the aerobic
2 ADP
metabolism of glucose from stored glycogen to
generate ATP. 2 ATP
• If the glycogen reserves are low, the muscle fiber can also Pyruvate
break down other substrates, such as fatty acids.
34 ADP
• All of the ATP now produced is used to power muscle 34 ATP
contraction.
CO2 To myofibrils to support
muscle contraction

Muscle Metabolism during Peak Activity
• During peak levels of activity, the demand for ATP is

enormous. Oxygen cannot diffuse into the fiber fast Lactate
enough for the mitochondria to meet that demand. Only
a third of the cell’s ATP needs can be met by the
mitochondria (not shown). Glucose Glycogen
• The rest of the ATP comes from glycolysis, and when this 2 ADP
produces pyruvate faster than the mitochondria can utilize ADP CP
2 ATP
it, the pyruvate builds up in the cytosol. This process is
called anaerobic metabolism because no oxygen is used. Pyruvate ATP
Creatine
• Under these conditions, pyruvate is converted to lactic
Lactate
acid, which dissociates into a lactate ion and a
hydrogen ion. To myofibrils to support
H+
muscle contraction
• The buildup of hydrogen ions increases fiber acidity,
which inhibits muscle contraction, leading to rapid fatigue.

• Muscle Fatigue
• When muscles can no longer perform a required
activity, they are fatigued
• Results of Muscle Fatigue
• Depletion of metabolic reserves
• Damage to sarcolemma and sarcoplasmic
reticulum
• Low pH (lactic acid)
• Muscle exhaustion and pain

• The Recovery Period

• The time required after exertion for muscles to
return to normal
• Oxygen becomes available
• Mitochondrial activity resumes

• Lactic Acid Removal and Recycling

• The Cori Cycle
• The removal and recycling of lactic acid by the liver
• Liver converts lactate to pyruvate
• Glucose is released to recharge muscle glycogen
reserves

• The Oxygen Debt

• After exercise or other exertion:
• The body needs more oxygen than usual to
normalize metabolic activities
• Resulting in heavy breathing
• Also called excess postexercise oxygen
consumption (EPOC)

• Heat Production and Loss

• Active muscles produce heat
• Up to 70 percent of muscle energy can be lost as
heat, raising body temperature

• Hormones and Muscle Metabolism

• Growth hormone
• Testosterone
• Thyroid hormones
• Epinephrine

10-7 Types of Muscle Fibers and Endurance
• Muscle Performance
• Force
• The maximum amount of tension produced
• Endurance
• The amount of time an activity can be sustained
• Force and endurance depend on:
• The types of muscle fibers
• Physical conditioning

• Three Major Types of Skeletal Muscle Fibers

1. Fast fibers
2. Slow fibers
3. Intermediate fibers

• Fast Fibers
• Contract very quickly
• Have large diameter, large glycogen reserves, few
mitochondria
• Have strong contractions, fatigue quickly

• Slow Fibers
• Are slow to contract, slow to fatigue
• Have small diameter, more mitochondria
• Have high oxygen supply
• Contain myoglobin (red pigment, binds oxygen)

• Intermediate Fibers
• Are mid-sized
• Have low myoglobin
• Have more capillaries than fast fibers, slower to
fatigue

Figure 10-21 Fast versus Slow Fibers.
Capillary
Slow fibers
Smaller diameter,
darker color due to
myoglobin; fatigue
resistant
LM × 170
Fast fibers
Larger diameter,
paler color;
easily fatigued
LM × 170 LM × 783

Table 10-2 Properties of Skeletal Muscle Fiber Types.

• Muscle Performance and the Distribution of

Muscle Fibers
• White muscles
• Mostly fast fibers
• Pale (e.g., chicken breast)
• Red muscles
• Mostly slow fibers
• Dark (e.g., chicken legs)
• Most human muscles
• Mixed fibers
• Pink

• Muscle Hypertrophy
• Muscle growth from heavy training
• Increases diameter of muscle fibers
• Increases number of myofibrils
• Increases mitochondria, glycogen reserves
• Muscle Atrophy
• Lack of muscle activity
• Reduces muscle size, tone, and power

• Physical Conditioning
• Improves both power and endurance
• Anaerobic activities (e.g., 50-meter dash,
weightlifting)
• Use fast fibers
• Fatigue quickly with strenuous activity
• Improved by:
• Frequent, brief, intensive workouts
• Causes hypertrophy

• Physical Conditioning
• Improves both power and endurance
• Aerobic activities (prolonged activity)
• Supported by mitochondria
• Require oxygen and nutrients
• Improves:
• Endurance by training fast fibers to be more like
intermediate fibers
• Cardiovascular performance

• Importance of Exercise
• What you don’t use, you lose
• Muscle tone indicates base activity in motor units
of skeletal muscles
• Muscles become flaccid when inactive for days or
weeks
• Muscle fibers break down proteins, become
smaller and weaker
• With prolonged inactivity, fibrous tissue may
replace muscle fibers

10-8 Cardiac Muscle Tissue
• Cardiac Muscle Tissue

• Cardiac muscle cells are striated and found only in
the heart
• Striations are similar to that of skeletal muscle
because the internal arrangement of myofilaments
is similar

• Structural Characteristics of Cardiac Muscle

Tissue
• Unlike skeletal muscle, cardiac muscle cells
(cardiocytes):
• Are small
• Have a single nucleus
• Have short, wide T tubules
• Have no triads
• Have SR with no terminal cisternae
• Are aerobic (high in myoglobin, mitochondria)
• Have intercalated discs

• Intercalated Discs
• Are specialized contact points between
cardiocytes
• Join cell membranes of adjacent cardiocytes (gap
junctions, desmosomes)
• Functions of intercalated discs
• Maintain structure
• Enhance molecular and electrical connections
• Conduct action potentials

• Intercalated Discs
• Coordination of cardiocytes
• Because intercalated discs link heart cells
mechanically, chemically, and electrically, the heart
functions like a single, fused mass of cells

Figure 10-22a Cardiac Muscle Tissue.
Cardiac
muscle cell
Intercalated
discs
Nucleus
Cardiac muscle tissue LM × 575
a A light micrograph of cardiac muscle tissue.

Figure 10-22b Cardiac Muscle Tissue.
Cardiac muscle
cell (intact)
Intercalated disc
(sectioned)
b A diagrammatic view of
cardiac muscle. Note
the striations and
intercalated discs.
Mitochondria
Nucleus
Intercalated Cardiac muscle cell

discs (sectioned)
Myofibrils

Figure 10-22c Cardiac Muscle Tissue.
Entrance to T tubule
Sarcolemma
Mitochondrion
Contact of sarcoplasmic
reticulum with
T tubule
Sarcoplasmic
Myofibrils reticulum
c Cardiac muscle tissue showing short, broad

T tubules and SR that lacks terminal cisternae.
• Functional Characteristics of Cardiac Muscle

Tissue
• Automaticity
• Contraction without neural stimulation
• Controlled by pacemaker cells
• Variable contraction tension
• Controlled by nervous system
• Extended contraction time
• Ten times as long as skeletal muscle
• Prevention of wave summation and tetanic
contractions by cell membranes
• Long refractory period
10-9 Smooth Muscle Tissue
• Smooth Muscle in Body Systems

• Forms around other tissues
• In integumentary system
• Arrector pili muscles cause “goose bumps”
• In blood vessels and airways
• Regulates blood pressure and airflow
• In reproductive and glandular systems
• Produces movements
• In digestive and urinary systems
• Forms sphincters
• Produces contractions

• Structural Characteristics of Smooth Muscle

Tissue
• Nonstriated tissue
• Different internal organization of actin and myosin
• Different functional characteristics

Figure 10-23a Smooth Muscle Tissue.
T
Circular
muscle layer
Longitudinal
muscle layer
Smooth muscle tissue LM × 100
a Many visceral organs contain several layers of smooth

muscle tissue oriented in different directions. Here, a
single sectional view shows smooth muscle cells in
both longitudinal (L) and transverse (T) sections.
Figure 10-23b Smooth Muscle Tissue.
Relaxed (sectional view)

Dense body
Actin Myosin
Relaxed (superficial view)
Adjacent smooth muscle cells are

Intermediate
bound together at dense bodies,
filaments (desmin)
transmitting the contractile forces
from cell to cell throughout the tissue.
Contracted
(superficial
view)
b A single relaxed smooth muscle cell is spindle shaped

and has no striations. Note the changes in cell shape as
contraction occurs.
• Characteristics of Smooth Muscle Cells

• Long, slender, and spindle shaped
• Have a single, central nucleus
• Have no T tubules, myofibrils, or sarcomeres
• Have no tendons or aponeuroses
• Have scattered myosin fibers
• Myosin fibers have more heads per thick filament
• Have thin filaments attached to dense bodies
• Dense bodies transmit contractions from cell to
cell

• Functional Characteristics of Smooth Muscle

Tissue
1. Excitation–contraction coupling
2. Length–tension relationships
3. Control of contractions
4. Smooth muscle tone

• Excitation–Contraction Coupling
• Free Ca2+ in cytoplasm triggers contraction
• Ca2+ binds with calmodulin
• In the sarcoplasm
• Activates myosin light chain kinase
• Enzyme breaks down ATP, initiates contraction

• Length–Tension Relationships
• Thick and thin filaments are scattered
• Resting length not related to tension development
• Functions over a wide range of lengths
(plasticity)

• Control of Contractions
• Multiunit smooth muscle cells
• Connected to motor neurons
• Visceral smooth muscle cells
• Not connected to motor neurons
• Rhythmic cycles of activity controlled by pacesetter
cells

• Smooth Muscle Tone

• Maintains normal levels of activity
• Modified by neural, hormonal, or chemical factors

Table 10-3 A Comparison of Skeletal, Cardiac, and Smooth Muscle Tissues.

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Chapter 10

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• Six Functions of Skeletal Muscle Tissue

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• Organization of Connective Tissues

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Skeletal Muscle (organ)

Epimysium Perimysium Endomysium Nerve

Muscle Muscle Blood

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Muscle Fascicle (bundle of fibers)

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Muscle Fiber (cell)

Capillary Myofibril Endomysium

Blood vessels Mitochondrion

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• Organization of Connective Tissues

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• Blood Vessels and Nerves

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Muscle fibers develop through

b A diagrammatic view and a

Mature muscle fiber

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Muscle fibers develop through

Mature muscle fiber

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Muscle fiber LM × 612

Sarcolemma Striations Nuclei

b A diagrammatic view and a

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Triad Sarcoplasmic T tubules

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H band Z line Titin

Zone of overlap M line Thin Thick

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Z line Zone of overlap M line

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Actinin Thin filaments Titin Thick filaments

Z line I band M line H band Zone of overlap

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Skeletal Muscle Myofibril

Surrounded by: Surrounded by: