10th E D I T I O N

MEDICAL PHYSIOLOGY

Guyton & Hall

Prof. Bi-Wen Peng,

Physiology,
Wuhan University
Peng Bi-Wen

Department of Physiology

email: pengbiwen@whu.edu.cn

cell phone: 1334 988 6616

 Unit II Membrane Physiology,
Nerve, and Muscle
6 Contraction of skeletal muscle

7 Excitation of skeletal Muscle

8 Contraction and excitation of smooth

muscle
May-June

Unit VII Respiration

37 Pulmonary Ventilation
38 Pulmonary Circulation
39 Physical Principles of Gas Exchange
40 Transport of Oxygen and Carbon
41 Regulation of Respiration
42 Respiratory insufficiency
 PPT show
Questions
Video show
Reviews
quiz
 Chapter 6:
Contraction of skeletal
muscle
 Part one
1. Anatomy of skeletal muscle
2. General mechanism of muscle contraction
3. Molecular mechanism of muscle contraction
4. Sliding filament theory
5. Motor unit and factors affecting force
production
 Muscle Tissue Types
Skeletal muscle40% body weight
Attached to bones
Nuclei multiple and peripherally located
Striated, Voluntary and involuntary (reflexes)
Smooth muscle
Walls of hollow organs, blood vessels, eye,
glands, skin
Single nucleus centrally located
Not striated, involuntary, gap junctions in
visceral smooth
Cardiac muscle
Heart
Single nucleus centrally located
Striated, involuntary, intercalated disks
VIDEO

Anatomy Muscle Types

 General Properties of Muscle
Contractility
Ability of a muscle to shorten with force
Excitability
Capacity of muscle to respond to a stimulus
Extensibility
Muscle can be stretched to its normal resting length
and beyond to a limited degree
Elasticity
Ability of muscle to recoil to original resting length
after stretched
 Skeletal Muscle
Human body contains over 400 skeletal muscles
40-50% of total body weight

Functions of skeletal muscle

Maintenance of posture
Respiration

Production of body heat

Constriction of organs and vessels

Basic Features of a Skeletal Muscle
Muscle attachments
Most skeletal muscles
run from one bone to
another
One bone will move
other bone remains
fixed
Origin less
movable attach-
ment
Insertion more
movable attach-
ment
Skeletal Muscle Structure
Composed of muscle cells
(fibers), connective tissue, blood
vessels, nerves
Fibers are long, cylindrical, and
multinucleated
Tend to be smaller diameter in
small muscles and larger in large
muscles. 1 mm- 4 cm in length
Develop from myoblasts;
numbers remain constant
Striated appearance
Nuclei are peripherally located
Structure of Skeletal Muscle:
Microstructure
 Muscle Fiber Anatomy

Sarcolemma - cell membrane

Surrounds the sarcoplasm (cytoplasm of fiber)
Contains many of the same organelles seen in other cells
An abundance of the oxygen-binding protein myoglobin

Punctuated by openings called the transverse tubules

(T-tubules)
Narrow tubes that extend into the sarcoplasm
Filled with extracellular fluid
 Myofibrils- cylindrical structures within muscle fiber
Threadlike strands within muscle fibers

Are bundles of protein filaments (=myofilaments)

Two types of myofilaments

Actin (thin filament), 3000 each myofibril, Troponin, Tropomyosin
Myosin (thick filament), 1500 each myofibril
At each end of the fiber, myofibrils are anchored to the inner surface of the sarcolemma
When myofibril shortens, muscle shortens (contracts)
Sarcoplasmic Reticulum (SR)
SR is an elaborate, smooth
endoplasmic reticulum
runs longitudinally and surrounds
each myofibril
Form chambers called terminal
cisternae on either side of the T-
tubules

A single T-tubule and the 2 terminal cisternae form a triad

SR stores Ca++ when muscle not contracting
When stimulated, calcium released into sarcoplasm
SR membrane has Ca++ pumps that function to pump Ca++ out of the
sarcoplasm back into the SR after contraction
 Part one
1. Anatomy of skeletal muscle
2. General mechanism of muscle contraction
3. Molecular mechanism of
muscle contraction
4. Sliding filament theory
5. Motor unit and factors affecting force
production
General mechanism of
muscle contraction

Initially, there is a local electrical event called end

plate potential
Later, it ignites an action potential that spreads in
all directions across the sarcolemma
muscle contraction

1. An action potential originating in the CNS reaches an alpha

motor neuron, which then transmits an action potential down its
own axon.
2. The action potential propagates by activating voltage-gated
sodium channels along the axon toward the neuromuscular
junction. When it reaches the junction, it causes a calcium ion
influx through the voltage-gated calcium channels.
muscle contraction

3. The Ca2+ influx causes vesicles containing the neurotransmitter acetylcholine to

fuse with the plasma membrane, releasing acetylcholine out into the extracellular
space between the motor neuron terminal and the neuromuscular junction of the
skeletal muscle fiber.

4. The acetylcholine diffuses across the synapse and binds to and activates nicotinic
acetylcholine receptors on the neuromuscular junction. Activation of the
nicotinic receptor opens its intrinsic sodium/potassium channel, causing sodium
to rush in and potassium to trickle out. Because the channel is more permeable to
sodium, the charge difference between internal and external surfaces of the
muscle fiber membrane becomes less negative, triggering an action potential.
muscle contraction

5. The action potential spreads through the muscle fiber's network

of T-tubules, depolarizing the inner portion of the muscle fiber.
6. The depolarization activates L-type voltage-dependent calcium
channels (dihydropyridine receptors) in the T tubule membrane,
which are in close proximity to calcium-release channels (ryanodine
receptors) in the adjacent sarcoplasmic reticulum.
muscle contraction

7. Activated voltage-gated calcium channels physically interact with calcium-

release channels to activate them, causing the sarcoplasmic reticulum to
release calcium.
8. The calcium binds to the troponin C present on the actin-containing thin
filaments of the myofibrils. The troponin then allosterically
modulates the tropomyosin. Under normal circumstances, the tropomyosin
sterically obstructs binding sites for myosin on the thin filament; once calcium
binds to the troponin C and causes an allosteric change in the troponin
protein, troponin T allows tropomyosin to move, unblocking the binding sites.
9. Myosin (which has ADP and inorganic phosphate bound to its nucleotide binding pocket
and is in a ready state) binds to the newly uncovered binding sites on the thin filament
(binding to the thin filament is very tightly coupled to the release of inorganic phosphate).
Myosin is now bound to actin in the strong binding state. The release of ADP and
inorganic phosphate are tightly coupled to the power stroke (actin acts as a cofactor in the
release of inorganic phosphate, expediting the release). This will pull the Z-bands towards
each other, thus shortening the sarcomere and the I-band.

10. ATP binds to myosin, allowing it to release actin and be in the weak binding state (a lack
of ATP makes this step impossible, resulting in the rigor state characteristic of rigor mortis).
The myosin then hydrolyzes the ATP and uses the energy to move into the "cocked back"
conformation. In general, evidence (predicted and in vivo) indicates that each skeletal
muscle myosin head moves 1012 nm each power stroke, however there is also evidence
(in vitro) of variations (smaller and larger) that appear specific to the myosin isoform.
muscle contraction

11 Steps 9 and 10 repeat as long as ATP is available and calcium

is freely bound within the thin filaments.
12 While the above steps are occurring, calcium is actively
pumped back into the sarcoplasmic reticulum. When
calcium is no longer present on the thin filament, the
tropomyosin changes conformation back to its previous state
so as to block the binding sites again. The myosin ceases
binding to the thin filament, and the contractions cease.
VIDEO

Muscular System, Neuromuscular

Junction
Part one

1. Anatomy of skeletal muscle

2. General mechanism of muscle contraction
3. Molecular mechanism of muscle contraction
4. Sliding filament theory
5. Motor unit and factors affecting force
production
MICROSCOPIC MUSCLE
1. Skeletal muscle consists of fibers (cells) covered by a sarcolemma and
contain sarcoplasm, nuclei, sarcoplasmic reticulum, and transverse
tubules.
These are involved in the pathway of the excitation-contraction
mechanism.
2. Each fiber contains thin and thick myofilaments.
The filaments are compartmentalized into sarcomeres
Micro Muscles (Cont.)

3. Thin filaments are composed of actin, tropomyosin, and

troponin; thick filaments consist of myosin.
a. Actin and myosin are the two contractile proteins in muscle.
b. Tropomyosin and troponin are muscle's regulatory proteins.
c. Projecting myosin heads are called cross bridges and contain actin-
and ATP-binding sites.
Sarcomere: Organization of Fibers
 Important:
3-dimensionally, thin
filaments are arranged
hexagonally around
thick filaments and the
cross-bridges project
from each thick
filament in all 6
directions towards the
surrounding thin
filamentseach thin
filament is surrounded
by 3 thick filaments.
Structure of Actin and Myosin
A sarcomere (Greek sar
x "flesh", meros "part") is
the basic unit of a
muscle.
Actin molecules are
bound to the Z line,
which forms the borders
of the sarcomere.
VIDEO

Action Potentials and Muscle Contraction

Questions

How does cross-bridge interaction between actin and

myosin bring about muscle contraction?
How does a muscle action potential trigger this
contractile process?
What is the source of the Ca2+ that physically
repositions troponin and tropomyosin to permit
cross-bridge binding?
Part one

1. Anatomy of skeletal muscle

2. General mechanism of muscle contraction
3. Molecular mechanism of muscle contraction
4. Sliding filament theory
5. Motor unit and factors affecting force
production
SLIDING FILAMENT THEORY

Definition:
When a muscle cell contracts, the thin filaments slide
past the thick filaments, and the sarcomere shortens.
This process comprised of several steps is called the
Sliding Filament Theory. It is also called the Walk
Along Theory or the Ratchet Theory.
 Physiology of Muscle Contraction

Steps Involved in Initiating Muscle Contraction

1. A neuron sends electrical signal (action potential) down
axon
2. Signal reaches ends of axon branches, stimulates release of
neurotransmitter acetylcholine (ACh)
3. ACh molecules cross synaptic cleft & bind with receptors in
sarcolemma
4. A muscle action potential travels along sarcolemma &
down transverse tubules
 Physiology of Muscle Contraction
(contd)
Events of muscle contraction
After the ATP has bound to the myosin head, the
binding of Myosin to Actin molecule takes place:
 Once the actin active sites are
uncovered, the myosin binds to it:
Power Stroke
POWER STROKE
 [Ca2+] in sacroplasm threshold

Abundent Ca2+ binds to troponin

Allosteric troponin

Allosteric tropomyosin and shifting

Uncovering binding site on actin

Cross bridge binds to actin

Head of cross brideg change its binding force

Cross bridge continues works Cross bridge twist towards to M line

as ATPase, hydrolyze ATP to Pull actin slide to center of myosin
provide energe
dissociation of cross bridge and
binding sitereposition

Cross bridge binds to next bindin gsite

Sacromere shorten
Sliding theory
 Physiology of Muscle Contraction (contd)

Sliding Filament Theory

1. Action potential travels to sarcoplasmic reticulum & releases
calcium ions into sarcoplasm
2. Calcium ions bind with troponin, moving aside tropomyosin
protein strands covering binding sites on actin filament
3. Myosin heads are charged with energy from breakdown of
ATP
4. Energy binds myosin heads to active receptor sites on actin
filament, making connections called cross-bridges
5. Ratcheting action (power stroke) occurs as myosin heads pull
sarcomere together, shortening the strand
6. Myosin heads bind more ATP, providing energy needed to
release hold on actin strand; process creates contractions
 Shortening of the
Muscle:

The thick and thin filaments

DO NOT shorten.
Contraction is accomplished
by the thin filaments from
opposite sides of each
sarcomere sliding closer
together or overlapping the
thick filaments further.
The H-zone becomes smaller
as the thin filaments approach
each other.
The I band becomes smaller as
the thin filaments further
overlap the thick filaments.
The width of the A band
remains unchanged as it
depends on the thick
filaments and the thick
filaments do not change
length.
 Mechanism of muscle contraction:
Width of sarcomere decreases from 2.2 to 2 or less
Length of thick and thin filament remain the same
Power stroke pull the Z discs towards the center
of the sarcomere
Results in app. Of the Z discs
Cyclic attachment-detachment-attachment of
myosin head to actin, till there is complete overlap
of thick & thin filaments
Decrease in I-band and H-band, A-band stays the
same
VIDEO

Muscular System, Sliding Filament

Theory (1) (2)
Part one

1. Anatomy of skeletal muscle

2. General mechanism of muscle contraction
3. Molecular mechanism of muscle contraction
4. Sliding filament theory
5. Motor unit and factors affecting force
production
 Neuromuscular junction
 Physiology of Muscle Contraction (contd)
Motor unit:
A motor unit consists of one somatic efferent (motor) neuron
and all of the muscle fibers (cells) that it innervates.
 Physiology of Muscle Contraction (contd)
Factors Affecting Force Production
Motor unit recruitment

Motor neuron: neuron responsible for initiating

motion
Motor unit: motor neuron & all fibers it controls

Few fibers = fine movement, less power; many

fibers = no fine movement, greater power
One muscle is typically composed of multiple
motor units
Body can control amount of force by varying
number & size of motor units recruited
Physiology of Muscle Contraction
(contd)
Factors Affecting Force Production
Cross-Sectional Area
As myofibrils become larger, muscles increase in cross-sectional
area & can generate more force
Fiber Arrangement
Pennate fiber arrangements generate more total force than
parallel fiber arrangements
Muscle Length
Shortened muscles have decreased ability to produce force
Muscles at resting length can produce greatest force
 Skeletal Muscle Fiber Types
Slow Twitch Fibers
Contract slowly but are resistant to fatigue
Rely on aerobic energy production
Used for long-duration activities (walking, jogging)
Fast Twitch Fibers
Contract rapidly & powerfully but fatigue quickly
Larger in diameter than slow twitch fibers due to having more
myofilaments
Rely on anaerobic energy production
Used for short-duration activities (sprinting, lifting)
 Skeletal Muscle Fiber Types
(contd)
Intermediate Fibers
Have characteristics of both slow twitch & fast twitch fibers
May adapt to bodys demands
Reservists waiting to be called up when & where need arises
Distribution of Fiber Types
Intermingled & genetically determined
Those with higher concentration of slow twitch fibers have
longer, leaner muscles & predisposition to endurance activities
Those with higher concentration of fast twitch fibers have larger,
thicker muscles & predisposition for sprinting or body building
 Slow and Fast Fibers
Slow-twitch or high-oxidative
Contract more slowly, smaller in diameter, better blood
supply, more mitochondria, more fatigue-resistant than
fast-twitch
Fast-twitch or low-oxidative
Respond rapidly to nervous stimulation, contain myosin
to break down ATP more rapidly, less blood supply, fewer
and smaller mitochondria than slow-twitch
Distribution of fast-twitch and slow twitch
Most muscles have both but varies for each muscle
Effects of exercise
Hypertrophies: Increases in muscle size
Atrophies: Decreases in muscle size
 PART TWO
1. Muscle Twitch
2. Graded Muscle Responses: Strength and
frequency
3. Muscle Tone
4. Preload and afterload
5. Isometric and Isotonic contraction
6. Muscle Fatigue
Muscle Twitch
A muscle twitch is the response of a muscle to a single
action potential of its motor neuron. The fibers
contract quickly and then relax.

Three phases:
Latent Period
Period of Contraction
Period of Relaxation

Myogram graphic recording of contractile activity

Muscle Twitch
Latent Period the first few ms after stimulation when
excitation-contraction is occurring
Period of Contraction cross bridges are active and the
muscle shortens if the tension is great enough to overcome
the load
Period of Relaxation Ca2+ is pumped back into SR and
muscle tension decreases to baseline level
Muscle Twitch
Twitch contraction of some muscles (extraocular) are
rapid and brief
others (gastrocnemius, soleus) are slower and longer
PART TWO

1. Muscle Twitch
2. Graded Muscle Responses: Strength and frequency
3. Muscle Tone
4. Preload and afterload
5. Isometric and Isotonic contraction
6. Muscle Fatigue
Graded Muscle Responses

Graded muscle responses are:

Variations in the degree or strength of muscle
contraction in response to demand
Required for proper control of skeletal
movement

Muscle contraction can be graded (varied) in two

ways:
By changing the Strength of the stimulus
By changing the Frequency of the stimulus
Stimulus Strength and Muscle Contraction

All-or-none law for muscle fibers

Contraction of equal force in
response to each action potential
Sub-threshold stimulus
Threshold stimulus
Stronger than threshold
Motor units
Single motor neuron and all
muscle fibers innervated
Graded for whole muscles
Strength of contractions range
from weak to strong depending on
stimulus strength
Muscle Response to Stimulation Frequency
A single stimulus results in a single contractile
response a muscle twitch (contracts and relaxes)
More frequent stimuli increases contractile force
wave summation - muscle is already partially
contracted when next stimulus arrives and
contractions are summed (refractory period applies)
Muscle Response to Stimulation Frequency

More rapidly delivered stimuli result in incomplete

tetanus sustained but quivering contraction
If stimuli are given quickly enough, complete
tetanus results smooth, sustained contraction with
no relaxation period
Muscle Response to Stronger Stimuli

Threshold stimulus the stimulus strength at

which the first observable muscle contraction
occurs
Beyond threshold, muscle contracts more
vigorously as stimulus strength is increased
Force of contraction is precisely controlled by
multiple motor unit summation
This phenomenon, called recruitment, brings
more and more muscle fibers into play
Stimulus Intensity and Muscle Tension

Below threshold no
muscle response
Above threshold increases
in voltage excite (recruit)
more (and larger) motor
units until maximal stimulus
is reached
Stimulus Frequency and Tension

Twitch Tetanic Contraction

Treppe: The Staircase Effect
Increased contraction tension in response to multiple
stimuli of the same strength. May be due to:
Increasing availability of Ca2+ in the sarcoplasm
Muscle enzyme systems become more efficient
and muscle pliability increases as muscle
contracts and liberates heat

Same intensity stimuli with

relaxation between
contractions the first few
contractions get stronger
and stronger
 PART TWO

1. Muscle Twitch
2. Graded Muscle Responses: Strength and frequency
3. Muscle Tone
4. Preload and afterload
5. Isometric and Isotonic contraction
6. Muscle Fatigue
Muscle Tone

Muscle tone:
The constant, slightly contracted state of all muscles -
does not produce active movements
Keeps the muscles firm and ready to respond to
stimulus
Helps stabilize joints and maintain posture
Due to spinal reflex activation of motor units in
response to stretch receptors in muscles and tendons
 PART TWO

1. Muscle Twitch
2. Graded Muscle Responses: Strength and frequency
3. Muscle Tone
4. Preload and afterload
5. Isometric and Isotonic contraction
6. Muscle Fatigue
1. Preload
1) Definition: the load encountered by the muscle
before it begins to contract
2) Effects: length-tension relationship
Muscle Length and Tension
Length-Tension Relationship in Skeletal Muscle
3) Mechanism
preload initial length
optimal initial maximal performance length

TF

AF AF

PF
2. Afterload

Definition: the load encountered by the muscle

after it begins to contract
Effects: force-velocity relationship

Force-Velocity Curve
AL LT Velocity

FT
Afterload (Force)
3) Mechanism
activity of ATPase and afterload

cross-bridge cycling

velocity of shortening

number of active cross-bridge force of contraction

 PART TWO

1. Muscle Twitch
2. Graded Muscle Responses: Strength and frequency
3. Muscle Tone
4. Preload and afterload
5. Isometric and Isotonic contraction
6. Muscle Fatigue
Contraction of Skeletal Muscle Fibers
The force exerted on an object by a contracting
muscle is called muscle tension, the opposing force or
weight of the object to be moved is called the load.
Two types of Muscle Contraction:
When muscle tension develops, but the load is not
moved (muscle does not shorten) the contraction is
called Isometric
If muscle tension overcomes (moves) the load and the
muscle shortens, the contraction is called Isotonic
Isometric & Isotonic Contractions

Isometric muscle contraction

develops tension without changing length
Isotonic muscle contraction
tension development while shortening = concentric
tension development while lengthening = eccentric
Isometric Contraction

Tension increases up to the muscles capacity, but

the muscle neither shortens nor lengthens
Occurs if the load is greater than the tension the
muscle is able to develop
The cross bridges generate force, but do not move
the thin filaments
Isometric Contractions
No change in overall muscle length

In isometric contractions, increasing muscle tension

(force) is measured
Isotonic Contraction
In isotonic contractions, the muscle changes length and
moves the load. Once sufficient tension has developed
to move the load, the tension remains relatively constant
through the rest of the contractile period.

Two types of isotonic contractions:

Concentric contractions the muscle shortens
and does work
Eccentric contractions the muscle contracts
as it lengthens
Isotonic Contraction
This illustrates a concentric isotonic contraction

In isotonic contractions, the amount of shortening

(distance in mm) is measured
 Resistance and Speed of Contraction
 PART TWO

1. Muscle Twitch
2. Graded Muscle Responses: Strength and frequency
3. Muscle Tone
4. Preload and afterload
5. Isometric and Isotonic contraction
6. Muscle Fatigue
Muscle Fatigue

Muscle fatigue the muscle is physiologically not

able to contract
Occurs when oxygen is limited and ATP production
fails to keep pace with ATP use
Lactic acid accumulation and ionic imbalances
may also contribute to muscle fatigue

When no ATP is available, contractures

(continuous contraction) may result because cross
bridges are unable to detach
Fatigue

Decreased capacity to work and reduced

efficiency of performance
Types:
Psychological
Depends on emotional state of individual
Muscular
Results from ATP depletion
Synaptic
Occurs in neuromuscular junction due to lack of
acetylcholine
Muscle Fatigue

Intense exercise produces rapid muscle fatigue (with

rapid recovery)
Na+-K+ pumps cannot restore ionic balances quickly
enough

Low-intensity exercise produces slow-developing

fatigue (with longer recovery period)
SR may be damaged, interfering with Ca2+ regulation
PART THREE

REVIEW
 REVIEWS & QUESTIONS
1. Whats the Properties of Muscle?
2. List the steps involved in initiating muscle
contraction.
3. Explain the sliding filament theory of skeletal
muscle?
4. What is a Neuromuscular Junction? Or, Whats
Excitation-Contraction Coupling?
5. Whats preload and afterload?
 1 Properties of Muscle
Contractility
Ability of a muscle to shorten with force
Excitability
Capacity of muscle to respond to a stimulus
Extensibility
Muscle can be stretched to its normal resting length
and beyond to a limited degree
Elasticity
Ability of muscle to recoil to original resting length
after stretched
2 Steps Involved in Initiating Muscle
Contraction

1. A neuron sends electrical signal (action potential)

down axon
2. Signal reaches ends of axon branches, stimulates
release of neurotransmitter acetylcholine (ACh)
3. ACh molecules cross synaptic cleft & bind with
receptors in sarcolemma
4. A muscle action potential travels along sarcolemma
& down transverse tubules
 3 What is a Neuromuscular
Junction?

1. Each skeletal muscle fiber connects to an axon from a nerve cell

called a motor neuron. The connection between the motor
neuron and muscle fiber is called a neuromuscular junction.
2. A neuromuscular junction (NMJ) is the synapse or junction of
the axon terminal of a motoneuron with the motor end plate,
3. The highly-excitable region of muscle fiber plasma membrane
responsible for initiation of action potentials across the muscle's
surface, ultimately causing the muscle to contract.
4. In vertebrates, the signal passes through the neuromuscular
junction via the neurotransmitter acetylcholine.
 3 Excitation Contraction Coupling
1. The train of physiological events that run from the
moment the brain decides to initiate a muscle
contraction to the actual contraction are termed
Excitation Contraction Coupling (EC Coupling).
2. The term "excitation" refers to the transmission of
a neural impulse (an Action Potential, AP) through
the peripheral nerves and eventually along the
sarcolemma and t-tubules.
3. "Contraction" obviously refers to the muscle
contraction and the biochemical events
responsible for the contraction. Coupling refers to
the linking of these two processes.
 4 Sliding Filament Theory
1. Action potential travels to sarcoplasmic reticulum & releases
calcium ions into sarcoplasm
2. Calcium ions bind with troponin, moving aside tropomyosin
protein strands covering binding sites on actin filament
3. Myosin heads are charged with energy from breakdown of ATP
4. Energy binds myosin heads to active receptor sites on actin
filament, making connections called cross-bridges
5. Ratcheting action (power stroke) occurs as myosin heads pull
sarcomere together, shortening the strand
6. Myosin heads bind more ATP, providing energy needed to
release hold on actin strand; process creates contractions
 At a very basic level each muscle fibre is made up
of smaller fibres called myofibrils. These contain
even smaller structures called actin and myosin
filaments. These filaments slide in and out
between each other to form a muscle
contractions, hence called the sliding filament
theory!

Sarcomere: The functional unit of the Myofibril, divided into I, A

and H bands.

The diagram shows part a myofibril called a sarcomere. This is

the smallest unit of skeletal muscle that can contract.
Sarcomeres repeat themselves over and over along the length
of the myofibril.
The diagram shows a fully contracted muscle with
lots of overlap between the actin and myosin.
Because the thin actin filaments have overlapped
there is a reduced potential for cross bridges to
form again. Therefore there will be low force
production from the muscle.
 5 Preload
Preload is the load given to the muscle before its
contraction.
Preload elongates the muscle thus enabling it to
contractin a prestretched state.
It determines the initial length of the muscle
before contraction.
Initial length is the length of the muscle fiber
before its contraction.
It is positively proportional to the preload.
Every muscle has its adequate preload or adequate
initial length.
 6 Effects of afterload

Afterload is the load given to the muscle

after the beginning of contraction.
When the afterload is zero, the muscle
contracts freely without load, the velocity of
contraction is at its maximum.