Excitable Tissues

Excitable Tissues

 Excitability is ability of a tissue to receive stimuli and respond to them by rapidly

changing their membrane potential.

 The stimuli can be mechanical, or electrical,

thermal.
 Two tissues in our body are said to be excitable:
[Link]
[Link]
 These tissues are excitable because they have high magnitude of
negative resting membrane potential and this RMP can undergo sudden
changes on stimulation resulting in the generation of what is known as
an action potential or nerve signal or impulse.
THE NERVE AS AN EXCITABLE TISSUE

The Nervous System Has Two Classes of Cells

These are:
Nerve cells (neurons) and
Glial cells (glia). Glial Cells Are Support Cells

•Glial cells far outnumber neurons—there are between 10 and 50

times more glia than neurons in the CNS of vertebrates.
•The name for these cells is derived from the Greek for glue, although
in actuality glia do not commonly hold nerve cells together.
•Rather, they surround the cell bodies, axons, and dendrites of
neurons.
• Glia are not directly involved in information processing, but they
are thought to have the following vital roles to play:
There are six types of glial cells:
1. Astrocyte –found in the CNS and for nutrient transfer to the
brain and spinal cord and form blood-brain barrier—that
prevents toxic substances in the blood from entering the brain
2. Some glial cells are scavengers –micorglia (CNS macrophages)
3. Ependymal cells –Line brain ventricles & facilitate
cerebrospinal fluid (CSF) conduction.
4. Oligodendrocytes –Form myelin sheath in the CNS.
5. Schwann Cells - Form myelin sheath in the PNS
6. Satellite cells –Found in the PNS but their function is not
known
NB: The first four are found in the CNS while the remaining two
are in the PNS.
Nerve Cells
Are the main signalling units of the NS: Generate AP and conduct
it.
A typical neuron has four morphologically defined
regions:
 Cell body
 Dendrites,
 Axon, and
 Presynaptic terminals.

Each of these regions has a distinct role in the generation and

communication of signals between nerve cells.
Fig. Structure of a Neuron
Classification of Neurons
Neurons may be classified in terms of their
1. Structure and
2. Function
1. Structural classification
Structurally neurons are classified into three:
1.1. Multipolar Neurons –have cell body, multiple
dendrites, and an axon. These are typical neurons
Multipolar neurons are found mainly in the CNS.
1. 2. Bipolar Neurons
The bipolar neurons have the cell body, one dendrite and an
axon. They are found in the retina and inner ear as well as in
the nasal mucous membrane
1. 3. The unipolar neurons
The unipolar neurons have a cell body and only one projection;
e.g., that found in the dorsal root ganglia of the grey matter of
spinal cord
2. Functional classification
 2.1. Sensory neurons- conduct information from periphery to
the CNS
 2.2. Motor neurons- conduct information from CNS to he
periphery
 2.3. Association neurons- integrate a sensory information and
provide feedback via the motor neurons.
Neuronal properties
 Excitability ( Generation of an action potential)
 Conductivity
 Membrane Potential and Resting Membrane
Potential

 Membrane potential is electrical environment of the cell.

 Or it is a charge separation between ICE & ECF due to
uneven distribution of ions across the cell membrane
 Resting membrane potential is a cell with its inside negativity
 This type of cell is called a polarized cell.
 Resting Potential Cont’d…
 Neurons are highly polarized (with a resting membrane
potential of about –90mV (RMP) due to at least 3factors:

Differential membrane permeability to K+ and Na+

through Na+-K+ leak channel (by far more permeable to
K+ than to Na+), which is by far the most important
The electrogenic nature of the Na+/K+ pump that pumps
3Na+ ions to the out side and 2K+ ions to the inside of
the cell.
The presence of intracellular non-diffusible anion
proteins.
 Changes in RMP with threshold stimulus allow generation of
action potentials and thus informative intercellular
communication occurs.
• Responsible for creation of MP
 Generation of Action Potential
If MP reaches threshold (-65 mV), →fast Na+ channels to open and
→ Na+ influx → depolarization.
Shortly Na+ channels become inactivated.
Opening of slow K+ channels → K+ efflux → repolarization.
This is known as an action potential
ECF
The successive stages of the action potential:
 Resting Stage
 Depolarization Stage.
 Repolarization Stage

1. Resting Stage: This is the resting membrane potential before

the action potential begins.
 The membrane is said to be "polarized" during this stage
because of the -90 millivolts negative membrane potential that
is present in the ICF of the cell.
 This means that the inside of membrane is about 90 times more
negative than the out side.
2. Depolarization Stage.

• The membrane suddenly becomes very permeable to Na+ after

sufficient stimulus is applied to the nerve, tremendous numbers
of Na+ diffuse/influx into the interior of the axon due to opening
of voltage- gated Na+ channels.
• The "polarized" state of -90 millivolts is neutralized and the
potential rising rapidly in the positive direction
• This stage of an AP is called depolarization. Also called rising
phase
• In large nerve fibers, the great excess of Na+ moving into the
fiber causes "overshoot" beyond the zero level
• In some smaller fibers, as well as in many CNS neurons, the
potential merely approaches the zero level and does not
overshoot to the positive state.
3. Repolarization Stage.
 Within a few 10,000ths of a second after the membrane becomes
highly permeable to Na+, Na channels begin to close and the
voltage-gated K+ channels open more than normal.
 Then, rapid efflux of K+ to the ECF re-establishes the normal
negative RMP.
This stage is called repolarization of the membrane.
• This is the falling phase.
• K+ channels are slow to open and slow to close. This causes the
RMP to take a brief below the usual RMP.
• This is called the undershoot and is an example of
hyperpolarization.
 90
-90
 Refractory Nature of Action Potential
There are two forms:
• Absolute refractory period (ARP)
• Relative refractory period (RRP)
Absolute refractory period (ARP)
• Interval b/n the opening of the Na+ channel activation gate
and its closing.
• 2nd AP can not be generated regardless of the strength of
stimulus applied.
• ARP begins at the start of up stroke (the activated Na+
channels start as fast as possible) and extends into the early
portion of downward stroke
• (Na+ channels are inactivated) .
• Na+ channel cannot be involved in another AP until the
inactivation gate has been reset.
 Relative Refractory Period
• New action potential can occur in an excitable fiber if the
stimulus is supra threshold
• The stimulus should be greater than normal b/c:
 There are still inactivated sodium channels
 More K+ channels than normal are still open.
• The RRP begins when the ARP ends.
• The action potential produced during this time has a lower up
stroke velocity and low overshoot potential than does the normal
AP
• Reason:
• Number of inactivated Na+ channels and
• Activated K+ channels during RRP .
 Properties of Action Potential
1. A Positive-Feedback Opens the Sodium Channels.

 First, as long as the membrane of the nerve fiber remains

undisturbed, no action potential occurs in the normal nerve.

 However, if any event causes enough initial rise in the

membrane potential from –90 millivolts toward the zero level,
the rising voltage itself causes many voltage-gated sodium
channels to begin opening.

 This allows rapid inflow of sodium ions, which causes a further

rise in the membrane potential, thus opening still more voltage-
gated sodium channels and allowing more streaming of sodium
ions to the interior of the fiber.
2. An action is generated with threshold stimulus
• This occurs when the number of Na+ ions entering the fiber
becomes greater than the number of K+ ions leaving the fiber.
• A sudden rise in membrane potential of 15 to 30 mV usually is
required.
3. Action potential has all or none property.
4. An action potential has no single direction of propagation.
• The action potential travels in all directions away from the
stimulus—even along all branches of a nerve fiber—until the
entire membrane has become depolarized.
 Action potential and graded potential
Graded potential
Has graded responses-Amplitude varies with condition of the
initiating event
Graded responses can be summated
Has no refractory period
Its amplitude decreases with distance
Duration varies
Can be depolarization or repolarization
 Initiated by environmental stimulus, NTs, or spontaneously
Action potential and Graded potential cont’d…
Action Potential
Amplitude is independent of the initiating event. It is all or none
Action potential can not be summated
Has refractory period
Its amplitude is not affected by distance
Duration is constant with a specific cell under constant condition
Is depolarization with an overshoot
Initiated by membrane depolarization.
 Action Potential Conduction
• If an AP is generated at the axon hillock, it will travel all the
way down to the synaptic knob.
• The manner in which it travels depends on whether the
neuron is myelinated or unmyelinated.
• Unmyelinated neurons undergo continuous conduction
• Myelinated neurons undergo saltatory conduction of an AP.
 Continuous Conduction
Occurs in unmyelinated
axons.

In this situation, the wave

of de- and repolarization
simply travels from one
patch of membrane to the
next adjacent
patch.
 Saltatory Conduction
• Occurs in myelinated
axons.
• Saltare is a Latin word
Saltatory Conduction
meaning “to leap.”
• Recall that the myelin
sheath is not completed.

• There exist myelin free

regions along the axon,
the nodes of Ranvier.
Rates of AP Conduction Depends Upon

Level of myelination
Faster in myrlinated than in unmyelinated
Size of nerve fiber
Faster in large sized than in smaller ones
Age
Slower in babies and in elderly
Maximum b/n the ages 5-15 years
 Synapses and synaptic transmission
• Synapse is an anatomical junction between pre-and post-
synaptic structures
• In autonomic NS, they are called ganglia (ganglion singular)
– Pre- synaptic structure is always a nerve
– Post- synaptic structure may be:
• A nerve
• Muscle
• Gland
• Skin
• The post- synaptic structures are collectively called effector
organs or simply effectors.
 Synaptic transmission cont’d…
Synaptic Transmission
• Begins with the stimulation of a neuron (Receptors) that
generates AP.
• Once stimulated, a neuron will communicate information
about the causative event.
– Such neurons are sensory neurons and they provide
information about both internal and external
environments.
 Synaptic transmission cont’d…
• Sensory (afferent) neurons will send information to the CNS
• Association neurons (interneurons) will integrate the
information
and send commands via motor (efferent) neurons which
synapse with effectors
– Thus, neurons need to be able to conduct information in 2
ways:
Through pre and post synaptic neurons, electrically.
Across the synapse, chemically.
 Synaptic transmission cont’d…
• One neuron will transmit information to another neuron or to a
muscle or gland cell by releasing chemicals called
neurotransmitters.
• The site of this chemical interplay is known as the synapse.
– An axon terminal (synaptic knob) will adjoin another cell, a
neuron, muscle fiber, skin, or gland cell.
– This is the site of transduction – the conversion of an
electrical signal into a chemical signal.
 Types of Junctions
There are types:
[Link] junction
[Link] junction
[Link] junction
Synaptic transmission cont’d…
 Mechanism of Chemical transmission

• An AP reaches the axon terminal

→open VG-Ca2+ channels.
Ca2+ rushes in, binds to
regulatory proteins & initiates
NT exocytosis.

NTs diffuse across the synaptic cleft and

then bind to receptors on the
postsynaptic membrane and initiate
some sort of response on the
postsynaptic cell.
 Effects of the Neurotransmitter
• Different neurons can contain different NTs.
• Different postsynaptic cells may contain different receptors.
– Thus, the effects of NT can vary.
• Some NTs cause cation channels to open, which results in a
graded depolarization.
• Some NTs cause anion channels to open, which results in a
graded hyperpolarization.
 Excitatory postsynaptic
potential (EPSP)
• A single synaptic interaction will
not create a graded depolarization

strong enough to induce the firing of

an AP.
– However, a graded
depolarization will bring the
neuronal MP closer to threshold.
– Thus, it’s often referred to as an
excitatory postsynaptic potential
( EPSP). This is due to cation
channel opening, leading to
cation influx
 Inhibitory postsynaptic potential (IPSP)

Graded hyperpolarizations
bring the
neuronal MP farther
away from threshold
and
thus are referred to as

inhibitory postsynaptic
potentials
(IPSP).
This is due to anion channel opening
leading to anion influx
 Summation
• One EPSP is usually not strong enough to cause an
AP.
• However, EPSPs may be summed to give rise to an
action potential.
There are two types of summation:
Temporal and spatial
[Link] summation
• This is when same presynaptic
neuron stimulates the postsynaptic
neuron multiple times in a brief period.
• EPSPs may be able to cause an AP.
2. Spatial summation
• Multiple presynaptic neurons all stimulate a
postsynaptic neuron resulting in a combination of
EPSPs which may yield an AP.
 Properties of synaptic transmission
 Unidirectional conduction
 Synaptic delay (0.5 -1.0m/s)
 Fatigue -↓in response of post-synaptic neurons after
repetitive stimulation by the pre-synaptic neurons
 Synaptic after discharge – persistence of out put signals
after the stoppage of pre -synaptic signals
 Synaptic potentiation (facilitation) –This is an ↑in post-
synaptic responses caused by previous post -synaptic
stimulation
 Neurotransmitter Removal

NTs are removed from the

synaptic cleft via:
Enzymatic
degradation
Diffusion
Reuptake
 Muscle Physiology

• Muscle is one of our 4 tissue types

• Found being combined with nerves, blood vessels, and
various connective tissues.
• Muscles are quite complex and are marvel of both biology
and physics.
 Muscle Functions

1. Produces Movement
2. Maintenance of posture
o Muscle contraction is constantly allowing us
to remain upright.
o The muscles of your neck keep your head up
right now.
o As you stand, your leg muscles keep you on
two feet.
3. Thermogenesis
o Generation of heat. Occurs via shivering –
an involuntary contraction of muscle.
4. Stabilization of joints
3 Types of Muscle Tissue
 Three Types of Muscle Tissue
 Skeletal Muscles
 Striated
 Multi-nucleatd
 Controlled by somatic nervous system (voluntary)
 Cardiac Muscle
 Striated
 Mono-nucleated
 Controlled by ANS (involuntary)
 Smooth Muscle
 Non-striated
 Mono-nucleated
 Controlled by ANS (involuntary)
 Characteristics of Muscle Tissue
1. Excitability- The ability to receive and respond to a
stimulus. The response is the generation of an AP that travels
along the plasma membrane of the muscle cell.

2. Contractility- The ability to shorten and become thicken

forcibly when adequately stimulated. This is the hallmark of
muscle tissue.

3. Extensibility - The ability to be stretched

4. Elasticity- The ability to recoil and resume original length

and size after being stretched or contracted.
 Skeletal Muscle Microanatomy
 Each skeletal muscle cell is known as a skeletal muscle fiber
because
• They are so long.
– Their diameter can be up to 100µm
– Their length can be as long as 30cm.
– They are so large because a single skeletal muscle cell results
from the fusion of hundreds of embryonic precursor cells
called myoblasts.
• A cell made from the fusion of many others is known as a
syncytium.
Muscle fiber
PM is known
As sarcolemm
muscle fiber
cytoplasm is
known as
sarcoplasm

Sarcolemma has invaginations that penetrate through the cell called

transverse tubules or T tubules.

Sarcoplasm has lots of mitochondria (why?), lots of glycogen

granules (to provide glucose for energy needs) as well as myofibrils
and sarcoplasmic reticuli.
 Sarcoplasmic Reticulum-SR

• SR is muscle cell version of the

smooth endoplasmic reticulum.
• Functions as a calcium storage
depot in muscle cells.
• Well developed in the skeletal
muscle
• Loose network of this
membrane bound organelle
surrounds all the myofibrils in
a muscle fiber.
 Myofibrils →ACTIN and MYOSIN
 Each muscle fiber contains rod-like structures called myofibrils
extend the length of the muscle cell.
 They are basically long bundles of protein structures called
myofilaments and their actions give muscle the ability to contract.
 The myofilaments are classified as thick filaments and thin
filaments.
 Myofilaments
 2 types of myofilaments (thick & thin) make up myofibrils.
 Thick myofilaments are made up of the protein myosin
A single myosin protein
resembles 2 golf clubs
whose shafts have been
twisted about one another

About 300 of these

myosin molecules are
joined together to form a
single thick filament
• Each thin filament is made up of 3 different types of protein: actin,
tropomyosin, and troponin.
– Each thin filament consists of a long helical double strand.
– This strand is a polymer that resembles a string of beads.
– Each “bead” is the globular protein actin.
– On each actin subunit, there is a myosin binding site.
– Loosely wrapped around the actin helix and covering the myosin binding
site is the filamentous protein, tropomyosin.
– Bound to both the actin and the tropomyosin are proteins collectively
known as troponin (A, I, C).
Note the relationship between the thin and thick filaments
 Myofibrils
• Each myofibril is made up 1000 of repeating individual units
known as sarcomeres.
• Each sarcomere is an ordered arrangement of thick and thin
filaments.
• Notice that it has:
– Regions of thin filaments by themselves (pinkish fibers)
– Region of thick filaments by themselves (purple fibers)
– Regions of thick filaments and thin filaments overlapping.
 Sarcomere
• The sarcomere is borderd by 2 protein structures
known as Z discs.
• The portion of the sarcomere which contains the thick
filament is known as the A band.
• A stands for anisotropic which is a fancy way of saying
that it appears dark under the microscope.
– The A band contains a zone of overlap (btwn thick & thin
filaments) and an H zone which contains only thick filaments
 The portion of the
sarcomere which
does not contain any
thick filament is
known as the I band.

The I band contains only

thin filament and is
light under the
microscope (it is
isotropic).

One I band is actually

part of 2 sarcomeres
at once. In the middle of the H zone is a structure
called the M line which functions to hold
the thick filaments to one another
 What Keeps the Myosin and Actin Filaments in Place?
 The side-by-side relationship between the myosin and actin filaments
is difficult to maintain.

 This is achieved by a large number of filamentous molecules of a

protein called titin.

 Each titin molecule has a molecular weight of about 3 million, which

makes it one of the largest protein molecules in the body.

 Also, because it is filamentous, it is very springy (Elastic).

 These springy titin molecules act as a framework that holds the

myosin and actin filaments in place so that the contractile machinery
of the sarcomere will work.
 Sliding Filaments
• All the sarcomeres in a fiber will contract together. This
contracts the fiber itself.
• The number of fibers contracting will determine the force of
the contraction of the whole muscle.
• The whole process of muscle contraction can be divided into 4
steps:
– Excitation
– Excitation-contraction coupling
– Contraction
– Relaxation
 Excitation
• All cells have a voltage difference across their plasma
membrane. This is the result of several things:
1. The ECF is very high in Na+ while the ICF is very high in
K+. The PM is impermeable to Na+ but slightly
permeable to K+. As a result, K+ is constantly leaking out
of the cell. In other words, positive charge is constantly
leaking out of the cell.
 Excitation

2. The Na+/K+ pump is constantly pumping 3 Na+ ions out and 2 K+ ions in for
every ATP used. Thus more positive charge is leaving than entering.
3. There are protein anions (i.e., negatively charged proteins) within the ICF
that cannot travel through the PM.
• What this adds up to is the fact that the inside of the cell is negative with respect
to the outside. The interior has less positive charge than the exterior.
• This charge separation is known as a membrane potential (MP).
• The value for MP in inactive muscle cells is typically btwn –80 and –90
millivolts.
• Cells that exhibit a MP are said to be polarized.
• MP can be changed by influx or efflux of charge.
 Excitation
• In general each muscle is
served by one nerve – a bundle
of axons carrying signals from
the spinal cord to the muscle.
• W/i the muscle, each axon will
go its own way and eventually
branch into multiple small
extensions called telodendria.
Each telodendrium ends in a
bulbous swelling known as the
synaptic end bulb.

The site of interaction btwn a neuron and any other cell is

known as a synapse. The synapse btwn a neuron and a
muscle is known as the neuromuscular junction.
 Excitation
The minute space between the synaptic end bulb and the
sarcolemma is known as the synaptic cleft.
There is a depression in the sarcolemma at the synaptic
cleft known as the motor end plate.
The synaptic end
bulb is filled with
vesicles that
contain the
neurotransmitter,
acetylcholine.
The motor end
plate is chock full
of acetylcholine
receptors.
 Excitation
1. A nerve signal will arrive at the synaptic end bulb and this will
cause the ACh-containing vesicles to undergo exocytosis.
2. ACh will diffuse across the synaptic cleft and bind to the ACh
receptors. These receptors are actually ligand-gated Na+
channels. The binding of ACh causes them to open.

3. Na+ will rush

into the cell,
making the
local cell
interior more
positive. This
is known as
depolarization
. It is a local
event!
 Excitation
• Adjacent to the motor end plate, the sarcolemma contains
voltage-gated ion channels.
• In order for these channels to open, the MP must depolarize
from its resting value of –90mV to approximately –50mV.
• This is the threshold. MP must become this much positive for
the voltage-gated channels to open.
• The degree of depolarization depends on how much Na+
influx occurred which in turn depends on how many Na+
channels were opened by binding Ach leading to the
generation of end plate potential (EPP).
 Excitation
• If the MP fails to depolarize to threshold, nothing will happen.

• The MP will soon return to normal and no muscle contraction

will occur.
• If the MP does reach threshold, 2 types of voltage-gated ion
channels will open:
– Fast Na+ channels
– Slow K+ channels
• If MP reaches threshold, fast Na+ channels open and Na+
rushes in causing the MP to depolarize to +30mV.
• The depolarization stops when the Na+ channels become
inactivated.
• At this point, slow K+ channels shall open & K+ efflux
occurs.
• This returns MP back to its resting level. This is
repolarization.
• This is known as an action potential.
 Excitation-Contraction Coupling

 It is a mechanism by which an action potential spreads in to the

skeletal muscle.
 The AP travels along the sarcolemma going in both directions
away from the motor end plate.
 Since T-tubules are simply invaginations of the sarcolemma,
the AP will spread down and through them as well.
• The T-tubular sarcolemma contains voltage sensitive proteins
that change their conformation in response to a significant Vm.
– These are physically linked to calcium channels in the SR membrane
– Upon Vm, the voltage sensors change their conformation. This
mechanically opens the Ca2+ channels in the SR membrane.
Excitation-Contraction Coupling
 Excitation-Contraction Coupling
The SR Ca2+ channels are only open briefly, but a large Ca2+
gradient exists so a large amount of calcium enters the sarcoplasm.

The Ca2+
interacts with the
2 regulatory
proteins of the
sarcomere so that
the 2 contractile
proteins can slide
& the sarcomere
can shorten.
 Contraction

• Normally, tropomyosin
obstructs the myosin
binding site on the G-actin
subunits.
• Calcium binds to the
troponin-C polypeptide of
the troponin triad.
• This changes the
conformation of troponin
which changes the
conformation of
tropomyosin which exposes
the myosin binding site on
actin.
 Contraction
• Once actin’s myosin binding site is exposed, myosin will
attach to it.
• Once myosin is bound to actin, the myosin head will
release the ADP and Pi which will cause it change
conformation.
• This results in the thin filament sliding along the thick
filament.
• Myosin then remains bound to actin until it binds to
another ATP.
• Myosin then hydrolyzes the new ATP and the cycle can
begin again.
 Mechanism of muscle contraction
Summary
(Excitation-contraction coupling)
When a muscle fibre membrane is depolarized, contraction of the
fibre follows.
The process by which depolarization initiates contraction is called
excitation contraction coupling.
It has several steps as follows:
1. Action potential initiated & propagated along the motor nerve
fibre and arrives at the end feet.
2. Opening of VG-Ca-channels and influx of Ca2+ to trigger the
release of Ach.
3. Ach released by Ca-dependent exocytosis and diffuse through
the synaptic cleft and binds to nicotinic receptors on post-
junctional membrane.
4. Opening of ligand gated Na-channels and influx of Na+ to
produce EPP.
5. Spread of depolarization through the sarcolemma
6. Spread of depolarization through the T-tubules
 Mechanism of muscle contraction
(Excitation-contraction coupling
7. Depolarization of T-tubules stimulate SR to release Ca 2+ into
sarcoplasm
8. Ca2+ binds to troponin-C
9. Ca2+ and troponin-C combination detaches troponin-I from the
active sites of actin
10. The detachment of troponin-I from actin displaces tropomyocin,
uncovering the active sites of actin filaments.
11. When the active site of actin is exposed, the heads of myosin
connect to them, making cross-bridges b/n myosin and actin.
12. The ATPase enzyme on the myosin heads hydrolyze ATP into ADP
+ -P plus energy.
13. The released energy causes the movement of the head (power
stroke) towards the centre.
14. The head of myosin is charged with a new molecule of ATP and
then detached from actin leading to relaxation.
 Relaxation
• Calcium pumps in the SR membrane work
constantly to get the calcium out of the
sarcoplasm and back into the SR. Ca2+ - pump
• They are unable to do this as long as the
muscle is still binding ACh.
• ACh is released by the motor neuron as long
as it keeps being stimulated.
• Note that ACh does not remain bound to the
AChR for very long.
• It quickly releases and either binds again or
more likely is hydrolyzed by the enzyme
acetylcholinesterase which exists as part of
the sarcolemma and free within the synaptic
cleft
 Relaxation
• When the muscle ceases
being stimulated, the calcium
pumps “win” and
sarcoplasmic [Ca2+] drops.
– Calcium stops being
available for troponin and
tropomyosin shifts back
into its inhibitory position.
• The muscle then returns back
to its original length via the
elasticity of the connective
tissue elements, plus the This animation shows
contraction of antagonistic another way to induce
muscles, and gravity. muscle relaxation.
 Mechanism of muscle relaxation
It has the following steps
1. Following muscle contraction, Ca2+ is re-up-taken back into SR
by Ca-pump, this requires ATP.
2. Decreased Ca2+ in the sarcoplasm→ Ca2+ detaches from
troponin-C →Tropomyosin covers the active sites of actin.
3. Head of myosin charged with ATP, and detached from actin
Therefore, muscle relaxation is an active process requiring
energy.
• Large amount of energy (ATP) is consumed during muscular
performance for the following activities:
1. To move the head of myosin (power stroke)
2. Active Ca2+ pump from sarcoplasm to SR
3. For Na-K-pump in the membrane
4. To remove the head of myosin from actin
• A motor unit is defined as a somatic Motor Units
motor neuron and all the skeletal
muscle fibers it innervates.
• When this neuron is stimulated, all
the muscle fibers it synapses upon
will be stimulated and will contract
as a unit
• The # of muscle fibers per motor
unit may be as high as several
hundred or as few as four.
– The smaller the motor unit, the
finer and more delicate the
movements.
– Extraocular muscles typically
have small motor units while the Notice that the muscle fibers of
large postural muscles have large a single unit are not clustered
motor units together but are spread out.
 Muscle Tone
• Some of the motor units with in particular muscle are
always active, even when the muscle is not
contracting.
– Their contractions do not produce enough tension to cause
movement, but they do tense and firm the muscle.
– This resting tension in a skeletal muscle is called tone.
– The identity of the motor units involved changes
constantly.

• Resting muscle tone stabilizes the position of bones

and joints.
Muscle Fiber Types

2 main types:
1. Slow fibers
2. Fast fibers
 Slow Fibers
• Contract slowly because its myosin ATPases work slowly.
• Depends on oxygen delivery and aerobic metabolism.
• Is fatigue resistant and has high endurance.
• Is thin in diameter – large amount of cytoplasm impedes O2 and
nutrient diffusion.
• Cannot develop high tension – small diameter means few
myofibrils.
• Has rich capillary supply and lots of mitochondria.
• Contains lots of the O2-storing protein, myoglobin which gives
it a red color.
• Uses lipids, CHO, and amino acids as substrates for its aerobic
metabolism.
• Best suited for endurance type activities.
• Red fibers, slow oxidative fibers, type I fibers.
 Fast Fibers
• So named because they can contract in 0.01 seconds or
less after stimulation.
• Fast fibers are large in diameter; they contain densely
packed myofibrils, large glycogen reserves, and
relatively few mitochondria.
• Able to develop a great deal of tension b/c they contain a
large number of sarcomeres.
• Use ATP in massive amounts. Supported by anaerobic
metabolism. Fatigue rapidly.
• fast fatigue (FF) fibers, fast glycolytic (FG) fibers,
white fibers.
• Best suited for short term, power activities.
 Smooth Muscle
• Involuntary, non-striated muscle tissue
• Occurs within almost every organ, forming sheets, bundles,
or sheaths around other tissues.
• Cardiovascular system:
– Smooth muscle in blood vessels regulates blood flow
through vital organs. Helps regulate blood pressure.
• Digestive systems:
– Rings of smooth muscle, called sphincters, regulate
movement along internal passageways.
– Smooth muscle lining the passageways alternates
contraction and relaxation to propel matter through the
alimentary canal.
 Smooth Muscle
• Integumentary system:
– Regulates blood flow to the superficial dermis & Allows for piloerection
• Respiratory system
– Alters the diameter of the airways and changes the resistance to airflow
• Urinary system
– Sphincters regulate the passage of urine
– Smooth muscle contractions move urine into and out of the urinary bladder
• Reproductive system
– Females
• Assists in the movement of the egg (and of sperm) through the female
reproductive tract
• Plays a large role in childbirth
– Males
• Allows for movement of sperm along the male reproductive tract.
• Allows for secretion of components of semen
• Allows for erection and ejaculation
 Smooth Muscle
• Smooth muscle cells:
– Are smaller: 5-10um in
diameter and 30-200um in
length
– Are uninucleate: contain one
centrally placed nucleus
– Lack any visible striations
– Lack T-tubules
– Have a scanty sarcoplasmic
reticulum

• Smooth muscle tissue is innervated by the autonomic nervous

system unlike skeletal muscle which is innervated by the
somatic nervous system (over which you have control)
Smooth Muscle
Contraction
• Myosin and actin are
present and crossbridge
formation powers
contraction, but the thick
and thin filaments do not
have the strict repeating
arrangement like that
found in skeletal muscle.
• Thin filaments are
attached to protein
structures called dense
bodies which attach to the
sarcolemma.
 Smooth Muscle
Contraction
• Begins with the opening of membrane
channels. Channels may be ligand-
gated (NTs, hormones, metabolites),
voltage-gated, or mechanically-gated
(stretch).
• Channels will allow significant calcium
entry from the ECF. Remember
smooth muscle has little SR.
• Calcium binds to a regulatory
molecule called calmodulin and
activates it.
• Activated calmodulin activates an
enzyme called Myosin Light Chain
Kinase.
 Smooth Muscle
Contraction
• Activated MLCK will add a
phosphate group to the
myosin of the thick filament.
This enables the myosin to
interact with actin.
– Tropomyosin is present
but not blocking actin’s
myosin binding sites
– Troponin is not present
• Contraction then ensues.
Smooth muscle relaxation:

Calcium is pumped out of the cell,

which decreases the amount of active
calmodulin which decreases the
amount of active MLCK which
decreases the number of crossbridges.

MLC phosphatase is required (removes

phosphate from the myosin)

Relaxation can occur subsequent to

contraction or at any time if anything
causes a decrease in the calcium
permeability of the smooth muscle cell.
Cardiac Muscle

• Striated, involuntary muscle

• Found in walls of the heart
• Consists of branching
chains of stocky muscle
cells. Uninucleate.
• Has sarcomeres & T-tubules
• Cardiac muscle cells are
joined by structures called Notice the branching
intercalated discs – which and the intercalated
consist of desmosomes and disc, indicated by the
gap junctions. blue arrow.