Lectures 6 and 7 hk468 PDF

Chapter 8
Skeletal Muscle: Structure and

Function
Copyright 2015 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of McGraw-Hill Education.
Chapter 8
Outline
Fiber Types
Structure of Skeletal
Muscle
Overview of
Neuromuscular
Biochemical and
Contractile
Junction
Characteristics of
Muscular Contraction
Skeletal Muscle
Overview of the Sliding
Functional
Filament/Swinging
Characteristics of
Lever Arm Model
Muscle Fiber Types
Energy for Contraction
Fiber Types and
Regulation of
Performance
Excitation-Contraction
Coupling
Exercise and Muscle
Fatigue
Exercise-Associated
Muscle Cramps
Muscle Actions
Speed of Muscle
Action and Relaxation
Force Regulation in
Muscle
Force-Velocity/ForcePower Relationships
Chapter 8
connective tissue called tendons. One end of the muscle is attached to a bone that does not move (origin),
and the opposite end is fixed to a bone (insertion) that
is moved during muscular contraction. A variety of different movements are possible, depending on the type
Skeletal muscle is composed

of several kinds
tissue.
Structure
ofofSkeletal
These include muscle cells (also called muscle fibers)
themselves, nerve tissue, blood, and various types of
connective tissue. Figure 8.1 displays the relationship
Structural Organization of Skeletal

Muscle
Tendon
Deep fascia
Artery
Nerve
Vein
Skeletal muscle
Epimysium
Perimysium
Nuclei
Perimysium
Artery
Vein
Nerve
Endomysium
Muscle fiber
Fascicle
Endomysium
(b) Fascicle
Endomysium
Muscle fiber
Nuclei
Myofibrils
(a) Muscle
Sarcoplasm
Sarcolemma
Satellite cell
(c) Muscle fiber
Figure Copyright
8.1 Structural
organization
of skeletal
muscle.
An entire
skeletalwithout
muscle
covered
by a ofconnective
tissue
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Education.
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No(a)
reproduction
or distribution
the is
prior
written consent
McGraw-Hill Education.
Muscle
Structure of Skeletal Muscle
Chapter 8
Connective Tissue Covering Skeletal

Muscle
Epimysium
Surrounds entire muscle
Perimysium
Surrounds bundles of muscle fibers
Fascicles
Endomysium
Surrounds individual muscle fibers
Basement membrane
Just below endomysium
Sarcolemma
Muscle cell membrane
Chapter 8
Satellite Cells
Play role in muscle growth
and repair
Increase number of nuclei
Myonuclear domain
Cytoplasm surrounding each nucleus
Each nucleus can support a limited
myonuclear domain
More nuclei allow for greater protein
synthesis
Important for adaptations to strength
training
Chapter 8
Microstructure of Muscle Fibers

Myofibrils
Contain contractile proteins
Actin (thin filament)
Myosin (thick filament)
Sarcomere
Includes Z line, M line, H zone, A band, I band
Sarcoplasmic reticulum
Storage sites for calcium
Terminal cisternae
Transverse tubules
Extend from sarcolemma to sarcoplasmic reticulum
are produced by alternating light and dark bands that
fore, to maintain a constant myonuclear domain, new

Structure
Muscle
nuclei (obtained from satellite
cells) of
areSkeletal
incorporated
into skeletal muscle fibers during growth (47, 49,
79). It follows that to maintain a stable myonuclear
Chapter
8 across the length of the fiber; the dark bands
appear
contain the primary muscle contractile proteins and

will be discussed in detail later.
Microstructure of Skeletal Muscle

Satellite cell
Skeletal
muscle fiber
Myofibril
Z line
Z line
Sarcomere
Filaments
of actin
(a)
(b)
H zone
I band
M line
A band
I band
Z line
A band
Myosin (thick)
filaments
Actin (thin)
filaments
(c)
Figure 8.2
TheCopyright
microstructure
of muscle.
Note
a skeletal
muscle
fiberwithout
contains
each consisting
2015 McGraw-Hill
Education. All
rightsthat
reserved.
No reproduction
or distribution
the priornumerous
written consent myofibrils,
of McGraw-Hill Education.
PAGES
StructureCONFIRMING
of Skeletal Muscle
Chapter 8
Structure of an Individual Skeletal

Muscle Fiber
Muscle
fiber
Nucleus
A band
I band
Z disc
Openings into
transverse tubules
Mitochondria
Sarcoplasmic
reticulum
Triad:
Terminal cisternae
Transverse tubule
Sarcolemma
Sarcoplasm
Myofibrils
Myofilaments
Figure 8.3
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2015
Education.
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reserved. Nofireproduction
distribution
the prior written consent of McGraw-Hill Education.
Structure
anMcGraw-Hill
individual
skeletal
ber. Seeortext
for without
details.
Chapter 8
Chapter 8
T-tubule network
* The T-system begins as invaginations of the membrane and is therefore
physically contiguous with the sarcolemma
* These invaginations extend transversely across the long axis of the
muscle fiber
Chapter 8
The sarcoplasmic reticulum
Neuromuscular Junction
Chapter 8
Neuromuscular Junction
Junction between motor neuron and muscle fiber
Motor unit
Motor neuron and all fibers it innervates
Motor end plate

Pocket formed around motor neuron by sarcolemma
Neuromuscular cleft
Short gap between neuron and muscle fiber
Acetylcholine is released from the motor neuron

Causes an end-plate potential (EPP)
Depolarization of muscle fiber
Chapter 8
The Neuromuscular Junction
Chapter 8
is always large enough to exceed threshold and is the

signal to begin the contractile process.
Acetylcholine is the neurotransmitter that stimNeuromuscular

ulates the muscle fiber to depolarize,
which is
the signal to start the contractile process.
The Neuromuscular Junction

IN SUMMARY
The human body contains more than 600 voluntary skeletal muscles, which constitute 40%
to 50% of the total body weight. Skeletal muscle performs three major functions: (1) force
production for locomotion and breathing,
(2) force production for postural support, and
(3) heat production during cold stress.
Motor nerve
fibers
Neuromuscular
junction
Muscle fibers
Motor nerve fiber
Myelin
Schwann cell
Synaptic knob
(a)
100 m
Synaptic vesicles
(containing ACh)
Basal lamina
Sarcolemma
Synaptic cleft
Nucleus
ACh receptor
Junctional folds
Mitochondria
Nucleus
Sarcoplasm
Myofilaments
(b)
Figure 8.4
The connecting point between a motor neuron and the muscle fiber is called the neuromuscular junction.
Junction
Chapter 8
The end-plate potential and the action potential
Chapter 8
Excitation-contraction coupling
Chapter 8
Excitation-contraction coupling
Chapter 8
Excitation-Contraction Coupling
Depolarization of motor end plate (excitation) is
coupled to muscular contraction
Action potential travels down transverse tubules and causes
release of Ca++ from SR
Ca++ binds to troponin and causes position change in
tropomyosin
Exposing active sites on actin
Strong binding state formed between actin and myosin

Contraction occurs
Chapter 8
The Sliding Filament Model

Also called the swinging lever-arm model
Muscle shortening occurs due to the movement of the
actin filament over the myosin filament
Formation of cross-bridges between actin and myosin
filaments
Power stroke
Reduction in the distance between Z lines of the

sarcomere
Chapter 8
CONFIRMING PAGES
Sarcomere Shortening During Muscle

Contraction
Sarcomere
Sarcomere
A band
A band
Z line
Actin
filaments
Z line
1 Relaxed
Z line
Z line
Myosin
filaments
2 Contracting
(a)
3 Fully contracted
(b)
Figure 8.5 When
a skeletal muscle contracts, (a) individual sarcomeres shorten as thick (myosin) and thin (actin)
Chapter 8
Sarcomere Shortening During Muscle

Contraction
http://videos.humankinetics.com/services/player/
bcpid1958213916001?bckey=AQ~~,AAAA0gHQGE~,UzAFL1pLzn5HKOtT4KHrsPjpUv67WEPE&bctid=
4142820261001
(a)
(b)
Chapter 8
Figure 8.5 When a skeletal muscle contracts, (a) individual sarcomeres shorten as thick (myosin) and thin (actin)
The Relationships Among Troponin,

Tropomyosin, Myosin, and Calcium
filaments slide past one another. (b) Photograph taken with an electron microscope that illustrates a shortened
sarcomere during muscle contraction (magnification is 23,0003).
Troponin
Calcium-binding site
Actin
Tropomyosin
Actin-binding
site
ATP-binding
site
Thin
filament
Myosin head
Myosin tail
Thick
filament
Figure 8.6 Proposed relationships among troponin, tropomyosin, myosin cross-bridges, and calcium. Note that when
Copyright 2015
McGraw-Hill Education.
All rights reserved.
No reproduction
or distribution
without
the priorand
writtencross-bridge
consent of McGraw-Hill
Education.
Ca11 binds to troponin,
tropomyosin
is removed
from
the active
sites on
actin
attachment
can occur.
Chapter 8
The Relationships Among Troponin,

Tropomyosin, Myosin, and Calcium
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d=4142821280001
Chapter 8
Molecular Steps in Cross-Bridge Cycling
Chapter 8
Molecular Steps in Cross-Bridge Cycling
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4161257754001
Fiber Types
Chapter 8
Characteristics of Muscle Fiber Types

Biochemical properties
Oxidative capacity
Number of capillaries, mitochondria, and amount of myoglobin
Type of myosin ATPase isoform

Speed of ATP degradation
Abundance of contractile protein in the muscle fiber

Actin and myosin
Fiber Types
Chapter 8

Contractile properties
Maximal force production
Force per unit of cross-sectional area
Speed of contraction (Vmax)

Myosin ATPase activity
Maximal power output

High force, fast fibers have high power output
Muscle fiber efficiency

Lower amount of ATP used to generate force
A Closer Look 8.2
Chapter 8
How Are Skeletal Muscle Fibers Typed?

Muscle biopsy
Small piece of muscle removed
May not be representative of entire body
Staining for type of myosin ATPase

isoform
Immunohistochemical staining
Selective antibody binds to unique myosin
proteins
Fiber types differentiated by color difference
Gel electrophoresis
Identify myosin isoforms specific to different
fiber types
A Closer Look 8.2
Chapter 8
Immunohistochemical Staining of
Skeletal Muscle
Blue = Type I fibers

Green = Type IIa fibers
Black = Type IIx fibers
Red = dystrophin (protein in sarcolemma)
Fiber Types
Chapter 8
Characteristics of Individual Fiber Types

Type I fibers
Slow-twitch fibers
Slow-oxidative fibers
Type IIa fibers

Intermediate fibers
Fast-oxidative glycolytic fibers
Type IIx fibers

Fast-twitch fibers
Fast-glycolytic fibers
Fiber Types
Chapter 8
CONFIRMING PAG
TABLE 8.1
Characteristics of Human Skeletal Muscle Fiber Types

fast fibers
Characteristic
Type IIx
Type IIa
slow fibers
Type I
Number of mitochondria
Resistance to fatigue
Predominant energy system
ATPase activity
Vmax (speed of shortening)
Efficiency
Specific tension
Low
Low
Anaerobic
Highest
Highest
Low
High
High/moderate
High/moderate
Combination
High
High
Moderate
High
High
High
Aerobic
Low
Low
High
Moderate
and contractile properties of human muscle fibers

several interesting facts have emerged. First, there
and can result in the conversion of fast fibers into
are no apparent sex or age differences in fiber disslow fibers (12, 35). A detailed discussion of exercisetribution (54). Second, the average sedentary man
induced changes in skeletal muscle fibers will be preor woman possesses approximately 50% slow fibers.
sented in Chap. 13.
Third, highly successful power athletes (e.g., sprintFinally, it is important to appreciate that while the
ers, etc.) typically possess a large percentage of fast
classification of skeletal muscle fibers into three distinct
fibers, whereas endurance athletes generally have a
Copyright 2015
McGraw-Hill
Education.the
All rights
reserved. No reproduction or
distribution
without the prior written
of McGraw-Hill
groups is a convenient
way
to group
functional
high
percentage
of consent
slow
fibers Education.
(22, 26, 73). Table
Fiber Types
Chapter 8
Comparison of Maximal Shortening

Velocities Between Fiber Types
Maximal shortening velocity
(muscle lengths second1)
4.0
3.5
3.0
2.5
2.0
1.5
1.0
0.5
0
Type I
Type IIa
Type IIx
Fiber type
Figure 8.12
Comparison of maximal shortening velocities between fiber types. Data are from reference 19.
Maximal specific
Chapter 8
Fiber Types
50
PAGES
ComparisonCONFIRMING
of Force
Production and
Power Output Between Fiber Types
Type I
Type IIa
Type IIx
40
30
Fiber type
Maximal specific force (kN/m2)
parison of maximal shortening velocier types.

Data are
from reference 19.
(a) Specific
force
Type I
Type IIa
Fiber type
Type IIx
(b) Maximal power
70
(W/t)
Maximal power (W/t)
) (19, 57). The mechanism to explain

sented earlier
(i.e., type I fibers possess a
60
TPase activity compared to fast fibers).
2
pe I fibers
produce a lower specific
50
ast fibers do (Fig. 8.13(a)) (19). The
1
this observation
is that type I fibers
40
in and myosin per cross-sectional area
I fibers do. This is important because
30
0
Type
IIa is diType IIx
orce generated Type
by a Imuscle
fiber
Type I
Type IIa
Type IIx
type
o the number of myosinFiber
cross-bridges
Fiber type
(i.e., the force generating state) at any
Figure 8.13 Comparison of (a) maximal specific force
mply (b)stated,
more cross-bridges
Maximalthe
power
production and (b) maximal power output between
e, the greater
the
force
production.
3
muscle fiber types. Data are from Canepari et al. (19).
fibers exert more force than slow fibers
ontain more myosin cross-bridges per
Fiber Types
Chapter 8
In Summary
Human skeletal muscle fiber types can be divided into three general
classes of fibers based on their biochemical and contractile properties
properties. Two categories of fast fibers exist: type IIx and type IIa.
One type of slow slow fiber exists: type I fibers.
The biochemical and contractile properties characteristic of all muscle
fiber types are summarized in table 8.1.
Fiber Types
Chapter 8
In Summary
Although classifying skeletal muscle fibers into three general groups
is a convenient system to study the properties of muscle fibers, it is
important to appreciate that human skeletal muscle fibers exhibit a
wide range of contractile and biochemical properties. That is, the
biochemical and contractile properties of type IIx, type IIa, and type I
fibers represent a continuum instead of three neat packages.
Fiber Types
Chapter 8
Fiber Types and Performance

Nonathletes
Have approximately 50% slow and 50% fast fibers
Power athletes
Sprinters
Higher percentage of fast fibers
Endurance athletes
Distance runners
Higher percentage of slow fibers
Fiber type is not the only variable that determines

success in an athletic event
esChapter 8
sum-
composition is not the only variable that determines

success in athletic events (22). In fact, success in athletic performance is due to a complex interaction of
psychological, biochemical, neurological, cardiopulmonary, and biomechanical factors (14, 15, 80).
Distribution of Fiber Type in Athletes
s into
em
t is
etal
is,
es of
ent a
s.
muscle
les, and
TABLE 8.2
Typical Muscle Fiber

Composition in Elite Athletes
Sport
Distance runners
Track sprinters
Nonathletes
% Slow
Fibers
(Type I)
% Fast Fibers
(Types IIx
and IIa)
7080
2530
4753
2030
7075
4753
Data from references 20 and 82.
Fiber Types
Chapter 8
Chapter 8
Single muscle fiber power from the world champion sprint runner compared with other
athletic and recreational populations from our laboratory.
Scott Trappe et al. J Appl Physiol 2015;118:1460-1466
2015 by
American
Physiological
Copyright
2015
McGraw-Hill
Education.Society
All rights reserved. No reproduction or distribution without the prior written consent of McGraw-Hill Education.
Fiber Types
Chapter 8
In Summary
Successful power athletes (e.g., sprinters) generally possess a large
percentage of fast muscle fibers and therefore a low percentage of
slow, type I fibers.
In contrast to power athletes, endurance athletes (e.g., marathoners)
typically possess a high percentage of slow muscle fibers and a low
percentage of fast fibers.
Although muscle fiber types are known to play a role in sport
performance, considerable variation exists among successful athletes
competing in the same sport
Muscle Actions
Chapter 8
Types of Muscle Action

Isometric
Muscle exerts force without changing length
Pulling against immovable object
Postural muscles
Dynamic (isotonic)
Concentric
Muscle shortens during force production
Eccentric
Muscle produces force but length increases
Associated with muscle fiber injury and soreness
Muscle Actions
Chapter 8
Summary of Muscle Actions

TABLE 8.3
Summary of the
Classifications of Exercise
and Muscle Action Types
Type of
Exercise
Muscle
Action
Dynamic
Concentric
Eccentric
Isometric
Static
Muscle
Length Change
Decreases
Increases
No change
Several types of muscle actions exist. For example, it is possible for skeletal muscle to generate force
without a large amount of muscle shortening. This can
Chapter 8
Muscle Actions
Isometric and Isotonic Muscle Actions
Speed of Muscle Action and Relaxation
Chapter 8

Muscle twitch
Contraction as the result of a single stimulus
Latent period
Lasting ~5 ms
Contraction
Tension is developed
40 ms
Relaxation
50 ms
Speed of shortening is greater in fast fibers

SR releases Ca++ at a faster rate
Higher ATPase activity
Chapter 8
Muscle Twitch
Time
40 ms
50 ms
Increasing force
of contraction
5 ms
Contraction
Relaxation
Latent
period
Stimulus
Figure 8.16 T
0 5
strength and
activate man
In contrast,
and more m
Figure 8.15
15
25
35
45
55
65
Time (milliseconds)
75
85
A recording of a simple twitch. Note the
95
Force Regulation in Muscle
Chapter 8

1) Types and number of motor units recruited
More motor units = greater force
Fast motor units = greater force
2) Initial muscle length

Ideal length for force generation
Increased cross-bridge formation
3) Nature of the neural stimulation of motor units

Frequency of stimulation
Simple twitch
Summation
Tetanus
CONFIRMING
PAGES
Force Regulation
in Muscle
Chapter 8
Relationship Between Stimulus Strength

and Force of Contraction
Increasing force
of contraction
Maximal response
Increasing stimulus strength

Figure
8.16 The relationship between increasing stimulus
Chapter 8
Length-Tension Relationships in
Skeletal Muscle
CONFIRMING PAGES
2.0 m
1.65 m
1.0
Relative tension
2.25 m
0.5
1.25 m
60
3.65 m
80
100
120
140
160
Percentage rest length
1.65 m
2.25 m
3.65 m
Less than
optimal length
Optimal length
Greater than
optimal length
Fewer cross-bridge
interactions = reduced
tension development
Maximal cross-bridge
interaction = maximal
tension development
Figure 8.17
No cross-bridge interaction
= no tension development
Length-tension relationships in skeletal muscle. Note that an optimal length of muscle exists, which will
produce maximal force when stimulated. Lengths that are above or below this optimal length result in a reduced
Copyright
2015
McGraw-Hill
Education. All rights reserved. No reproduction or distribution without the prior written consent of McGraw-Hill Education.
amount
of force
when
stimulated.
be closely replicated in the laboratoryForce

if aRegulation
seriesin of
Muscle
stimulations are applied to the muscle. The recording
Simple
Twitch, Summation, and Tetanus
pictured in Fig. 8.18 represents what occurs when successive stimuli are applied to the muscle. The first few
Chapter 8
Force
Tetanus
Simple
twitches
Summation
Stimuli
Figure 8.18
Recording showing the change from simple

twitches to summation, and finally tetanus. Peaks to
the left represent simple twitches, while increasing the
Chapter 8
Force-Velocity / Power-Velocity Relationships
Force-Velocity Relationship
At any absolute force the speed of movement is
greater in muscle with higher percent of fast-twitch
fibers
The maximum velocity of shortening is greatest at the
lowest force
True for both slow and fast fibers
Chapter 8
Force-Power Relationship
At any given velocity of movement, the power
generated is greater in a muscle with a higher percent
of fast-twitch fibers
The peak power increases with velocity up to
movement speed of 200300 degreessecond1
Power decreases beyond this velocity because force decreases
with increasing movement speed
- Chaptertween
muscular force and the
speed / of
movement.
Force-Velocity
Power-Velocity
Relationships
8
The relationship between speed of movement and
r Muscle
muscularForce-Velocity
force is shown in Fig.Relationships
8.19.
e
n
80% fast fibers
100
.
d
e
h
f
% Maximal force
80% slow fibers
Velocity of movement
Figure 8.19 Muscle force-velocity relationships. Note that
at any given speed of movement, muscle groups with a

Chapter 8
Muscle Force-Power Relationships
cle,
ater in
fast
ssess
g is
gainst the
ment
). This
bers.
he forceis of the
yosin. Remuscle is
Power output
examina-
80% fast fibers

80% slow fibers
0
Figure 8.20
100
200
300
400
Velocity of movement (degrees/second)
Muscle power-velocity relationships. In gen-
Chapter 8
In Summary
The amount of force generated during muscular contraction is
dependent on the following factors: (1) types and number of motor
units recruited, (2) the initial muscle length, and (3) the nature of the
motor units neural stimulation.
The addition of muscle twitches is termed summation. When the
frequency of neural stimulation to a motor unit is increased, individual
contractions are fused in a sustained contraction called tetanus.
The peak force generated by muscle decreases as the speed of
movement increases. However, in general, the amount of power
generated by a muscle group increases as a function of movement
velocity.
Chapter 8
Myo-mechanical Analysis of Isolated Skeletal Muscle
http://www.jove.com/video/2582/myo-mechanicalanalysis-of-isolated-skeletal-muscle
Chapter 8
Assessment of Neuromuscular
Function Using Percutaneous
Electrical Nerve Stimulation
http://www.jove.com/video/52974/assessmentneuromuscular-function-using-percutaneous-electricalnerve

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Chapter 8

Skeletal Muscle: Structure and

Skeletal muscle is composed

Structural Organization of Skeletal

Structure of Skeletal Muscle

Connective Tissue Covering Skeletal

Structure of Skeletal Muscle

Structure of Skeletal Muscle

Microstructure of Muscle Fibers

are produced by alternating light and dark bands that

fore, to maintain a constant myonuclear domain, new

contain the primary muscle contractile proteins and

Microstructure of Skeletal Muscle

Structure of an Individual Skeletal

The sarcoplasmic reticulum

Motor end plate

Acetylcholine is released from the motor neuron

The Neuromuscular Junction

is always large enough to exceed threshold and is the

Acetylcholine is the neurotransmitter that stimNeuromuscular

The Neuromuscular Junction

The end-plate potential and the action potential

Strong binding state formed between actin and myosin

The Sliding Filament Model

Reduction in the distance between Z lines of the

Sarcomere Shortening During Muscle

Sarcomere Shortening During Muscle

The Relationships Among Troponin,

The Relationships Among Troponin,

Molecular Steps in Cross-Bridge Cycling

Molecular Steps in Cross-Bridge Cycling

Characteristics of Muscle Fiber Types

Type of myosin ATPase isoform

Abundance of contractile protein in the muscle fiber

Characteristics of Muscle Fiber Types

Speed of contraction (Vmax)

Maximal power output

Muscle fiber efficiency

A Closer Look 8.2

How Are Skeletal Muscle Fibers Typed?

Staining for type of myosin ATPase

A Closer Look 8.2

Blue = Type I fibers

Characteristics of Individual Fiber Types

Type IIa fibers

Type IIx fibers

Characteristics of Human Skeletal Muscle Fiber Types

and contractile properties of human muscle fibers

Comparison of Maximal Shortening

Maximal specific force (kN/m2)

parison of maximal shortening velocier types.

(b) Maximal power

Maximal power (W/t)

) (19, 57). The mechanism to explain

Fiber Types and Performance

Fiber type is not the only variable that determines

composition is not the only variable that determines

Distribution of Fiber Type in Athletes