34

Excitable Tissues

Physiology Of Excitable Tissues

Excitable tissues
All tissues in our body are living cells and can maintain resting
membrane potential. Excitability, however, is shown only by specialized
cells, nerve and muscle. An excitable cell is able to respond to a stimulus
and generare electrical changes.

Neurons
Nerve cells, called neurons, are responsible for conducting impulses
from one part of the body to another. They are the structural and functional
units of the nervous system.

A neuron consists of 3 portions.

1) cell body. 2) dendrites. 3) axon.
Parts Character
The cell Contains a well-defined nucleus surrounded by a cytoplasm.
Within the cytoplasm are typical organelles such as mitochondria,
body lysosomes and golgi complex.
Dendritie Are branched, extensions of the cell body.
s Their function is to conduct impulses toward the cell body.
Axon It is a single, long, thin process that conduct impulses way from
the cell body to another neuron.
This axon formed of initial sagment and terminal branches called
synaptic knobes which contain synaptic vesicles.
When the impulse reaches the synaptic terminals leading to release of
neurotransmitter which excite or inhibit nearby neurons
It may be myelinated or unmyelinated.
This myelin contain gaps called nodes of Ranvier.
This myelin is protective and electrical insulator, it also incease the
speed of transmission of nerve impulse.

Properties of nerves
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Excitable Tissues

1- Excitability: it the ability of nerve to respond to stimulus.

2- Conductivity: it the ability of nerve to conduct nerve impulse.
3- Infatiguibility: nerves not liable to fatigue by repeated stimulation.
4- All or none law:
If a single nerve fiber is stimulated with a
threshold or suprathershold stimulus, an action
potential will result, which always have the
same characteristic amplitude regardless the
strength of stimulation.
Subthreshold stimulus ; no action potential.
Threshold stimulus; action potential.
Suprathreshold stimulus; same action potential.

Stimulus
Def: It is a change in the surrounding environment.
Types: electrical, thermal, chemical or mechanical.
Electrical stimuli are used to study the function of nerve or muscle. It is
preferred because intensity and duration can be easily controlled.

Effects of stimulation of the nerve

A given stimulus will cause its effect by altering the
permeability to one or more ions. The involved ions will then
diffuse into or out of the cell according to their concentration
and electrical gradients, causing a change in the membrane
potential.

Depolarization occurs when the membrane potential becomes less negative,

moving toward zero. Depolarization makes the neuron more excitable.

Hyperpolarization occurs when the membrane potential becomes more

negative, moving away from zero. Hyperpolarization makes the neuron less
excitable.

Once the stimulus has been removed, the membrane potential returns to its
resting state. Following depolarization, the membrane is said to undergo
repolarization, returning to its resting potential.

Resting membrane potential

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Definition:
It is the difference in potential under resting conditions between the
outer surface and inner surface of the membranes of excitable tissues with the
inside of the membrane negatively charged with respect to the outside.
The positivity of the outer surface is due to Na+ ions; while the negativity of
inner surface is due to intracellular protein anions.

RMP is normally 70 mv in nerves and 90 mv in muscles

Causes of resting membrane potential

1) Selective permeability of the nerve membrane:
* permeability of membrane to K+ is more than that Na+ at rest.
* Passive K+ efflux is more than passive Na+ influx.
* K+ efflux is not accompanied by equal efllux of anions (-ve).
* The net result is that the membrane is maintained in a polarized state with
the outside positive relative to the inside.

2) Sodium-potassium pump:
*The Na+K+ pump also plays a vital role in
this process.
*For each molecule of ATP expended, three
Na+ ions are pumped out of the cell into the
ECF and two K+ ions are pumped into the
cell into the ICF.
*The result is the unequal transport of
positively charged ions across the membrane
such that the outside of the cell becomes more
positive compared to its inside; in other
words, the inside of the cell is more negative
compared to the outside.

Action potential
Definition:
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It is a sudden change in the membrane potential of an excitable cells of the

nerve in response to stimulus.

Phases
1) Depolarization stage
At the beginning of A.P, depolarization is slow for 15 mv till it reaches the
firing threshold then the rate of depolarization increases markedly.
2) Then the membrane potential overshoots and becomes + 35 mv.
3) Repolarization stage
At the end of depolarization, the change in membrane potential then
reverses and falls rapidly towards the resting level.

Ionic basis
1- At the beginning of action potential,
voltage gated Na channeles start to
open, so the membrane potential reach
to 15 mv (firing level).
2- At the firing level, of the action
potential, sodium channels open
(activation state) increasing
+
permeability to Na several hundreds
fold. Na+ rushes into the membrane.
3- Positive feedback cycle develops:
i) Depolarization opens Na+
channels.
ii) Na+ diffuses into the cell due to
increased Na+permeability.
iii) Addition of +ve charges into the
cell further depolarizes the
membrane.
iv) This in turn produces a still greater
increase in Na+ permeability which inturn causes and so on.
4- In the repolarization phase what causes
the membrane potential to return so rapidly
to its resting level
- It is due to opening of K+ channels &
closure of Na channel.
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5- Due to to delayed closure of K+ channels, the membrane potential

undershoot resulting in hyperpolarization.

6- The membrane potential returns to the resting state by activation of

sodium potassium pump.

Firing Level

Excitability changes during action potential

1) Absolute Refractory Period

During depolarization and early repolarization, there is no response to any
stimulus whtever its strength. The excitability is zero.
2) Relative refractory period
During the lower 2/3 of repolarization, a stimulus stronger than threshold
can stimulate the nerve.

Propagation of the nerve impulse

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a) Conduction in unmyelinated fibers

The (+) charges in the region
of the action potential are
attracted to the negative
charges in the immediately
adjacent region of the axonal
membrane.
This current flow depolarizes
the new region, causing an
increase in the permeability of
the membrane to Na+ ions. The subsequent influx of Na+ ions further
depolarizes the membrane so that it reaches threshold and a new action
potential is generated in this region.
At the same time, the original site of action potential generation
repolarizes due to the efflux of K+ ions.

b) Conduction in myelinated fibers

Action potentials
cannot be generated in
the regions covered
with myelin. Instead,
they occur only at
nodes of Ranvier.
The flow of current
from an active node
skips down the axon
to the adjacent node to
cause depolarization
and generation of a new
action potential.
This transmission of the impulse from node to node is
referred to as saltatory conduction. (The term saltatory
means jumping ).

Saltatory conduction is of value for two reasons

First it increases the velocity of nerve transmission.
Second it conserves energy, since depolarization occurs only at the node.

Factors Affecting Conduction Velocity

1. Types of nerve fibers :
I- Type A (alpha, Beta, Gamma, Delta).
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It has the largest diameter (3-20 microns) and the highest Speeds of
conduction (15-120 meters/sec).
II- Type B
It has a diameters (1.3-3 microns) and speed of conduction (3-15
metes/second).
III- Type C
It has the diameter of 0.3-1.3 microns and speed of conduction 0.5-3
meters/sec.(smallest diameter & slowest conduction).

A & 8 fibers are myelinated, while C fibers are unmyelinated.

The greater the nerve diameter, the greater is the speed of conduction. Therefore,
conduction velocity of the myelinated nerve fiber is faster than the unmyelinated
one.
Normal propagation along an non-myelinated axon is typically in the range of 1-2
m/sec. In myelinated axon, this ranges from 15 to 150 m/sec.

2. Body temperature :
One degree fall in body temperature decreases nerve conduction
velocity by 3%. At 3 - 7 C, nerve conduction is completely blocked. This
physiologic observation is applied in many beneficial surgical procedures
(cryosurgery).

Local potential

DeFinition
The local potential is the depolarization of a cell below threshold. After
the cell is sufficiently depolarized (and reaches threshold), it fires an action
potential down the axon.
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Characteristic features of the local potential

Its magnitude is proportionate to the strength of the stimulus = graded
response (amplitude-modulated).
It is localized to the area of stimulation = not propagated.
it could be summated.
Local potentials could be induced by sub-threshold stimulation of the
excitable membranes of the nerve and muscle fibers.
Local potentials are important in short distance signaling function.

Neuromuscular Transmission
Motor end plate (neuromuscular junction) MEP
It is the area of contact between motor nerve fiber and muscle fibers.

Mechanism of neuromuscular transmission

a) Development of end-plate potential
-Action potentials in the motor neuron cause release of the
neurotransmitter acetylcholine.
-Binding of acetylcholine to its receptors on the muscle fiber causes an
increase in the permeability to Na+ and K+ ions.
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-The ensuing depolarization generates an action potential that travels

along the surface of the muscle fiber in either direction that is referred to
as a propagated action potential.
-This action potential elicits the intracellular events that lead to muscle
contraction.

b) Destruction of acetylcholine by choline esterase

The released acetylcholine is rapidly hydrolysed by choline-esterase
enzyme. This prevents re-excitation of the muscle.

Properties of NM transmission
a) Unidirectional: from the nerve to the muscle.
b) NM delay: 0.5 msec passed from the action potential in the motor nerve
terminals to the start of the muscle contraction (Ca influx, release of ach,
binding of acetylcholine to the receptors and deveelopment of muscle action
potential).
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Excitable Tissues

c) NM fatigue: decrease in the NM transmission upon repeated stimulation

of the nerve terminals due to depletion of acetylcholine.

d) Drugs affecting neuro-muscular transmission

A) Drugs block transmission
1) Curare (d-tubocuraine) acting as competitive antagonist of acetylcholine
at motor end plate.
2) Botulinum toxins: by inhibiting the release of acetylcholine.

B) Drugs stimulate transmission

1) Acetylholine-like drugs e.g. metacholine, carbachol and nicotine.
2) Anticholine esterase drugs e.g. prostigmine, eserine and DFP.

Anticholine-esterases
Short acting
- eserine (physostigmine). - prostigmine (neostigmine).
These drugs increase the availability of acetylcholine by preventing its rapid
breakdown by the choinestrase.

Long acting
- Parathion (isecticides) - Di-isopropyl fluorophosphates (war poison).
They produce irriversible inhibition of cholinestrae enzyme, resulting in
persistnt depolarization and muscle paralysis. Death may follow due to
failure of respiration.

Myasthenia gravis
It is characterized by marked weakness and easy fatigueability of muscles. It
may be due to:
Autoimmune disease caused by the formation of circulating antibodies
destroying the acetylcholine receptors. It is treated by anti-cholinestrases

Skeletal muscle
Skeletal muscles have the ability to use chemical energy to produce
force and movement.
It is called striated muscle because its fibers exhibit alternating light and dark
bands called striations.
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Structure of skeletal muscle

1) Muscle fibers
A single skeletal muscle consists of
many multinucleated muscle Cells called
muscle fibers.
2) Myofibrils
Each muscle fiber is composed of
longitudinally parallel structures called
myofibrils. It is divided into functional
units called sarcomere.
3) Myofilaments
The myofibril contains thick filaments
and thin filaments.
4) Striations
The myofibrils are crossed from side to
side by numerous dark and light bands
called striations.
The dark bands are called A bands, and
the light bands are called I bands.
Sarcomeres;
are separated from one another by
vertical Z lines.
In the center of A band there is H zone
corresponds to the space between the
ends of the two thin filaments. It also
marks the region which contains no
cross bridges.

There are two types of filaments: thick and thin

a) The thick filament; formed of myosin molecules
b) The thin filament
The thin filament is also complex: actin, tropomyosin and troponin.
The thick - it formed of 2 globular heads projecting at one end and the tail.
filament - Each head of myosin molecule contains an actin binding site and
myosin ATPase site (where ATP bind).
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(myosin) Head of myosin can be called cross bridge.

The thin Actin, tropomyosin and troponin.

filament
Actin There are active sites on the actin filaments with which the cross
bridges of myosin interact to cause muscle contraction.
Troponin which is formed of 3 subunits:
o Troponin T: binds to tropomyosin.
o Troponin I: binds with actin & prevents the filaments from sliding
when at rest
o Troponin C: has two regulatory bindings sites for Ca.
Tropomyosin In resting state, it covers the active sites of the actin.
So that interaction cannot occur between the actin and myosin to
cause contraction (relaxing protein).
Troponin It is a complex of three proteins. Troponin I has strong affinity of
actin, Troponin T for tropomyosin, and Troponin C for Ca++.
The sarcoplamic It collect (during muscle relaxation) and release Ca ++ around the
reticulum myofibril. (during muscle contraction)
Transverse - The cell membrane invaginates to form transverse tubules.
tubules: T.T - They open to the outside of the cell.
- T.T serve to conduct electrical excitation (action potential) deep
into the muscle fiber from the surface membrane.

Excitation-Contraction Coupling

Definition
It is the process by which depolarization of the muscle fiber initiates
contraction. The link between these events is the calcium ion.
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Mechanism
1) When the nerve impulse reached the neuro-muscular junction, thus acetyl
choline is released by exocytosis.
2) Acetyl choline combines to specific nicotinic receptors on muscle.
3) The combination between Ach and the receptors results in increase in ionic
permeability of the muscle membrane, give rise to action potential.
4) Action potential travel over the muscle fiber membrane.
5) Inward spread of depolarization along T tubules Release of Ca ++ from
the Sarcoplasmic Reticulum into cytoplasm.
6) Binding of Ca++ to troponin C, as a result, the troponinactin linkage is
weakened, allowing the tropomyosin to be repositioned such that the myosin-
binding sites are uncovered. The myosin crossbridge now binds to the actin,
causing the energy previously stored within the myosin to be discharged and
the crossbridge to swivel inward toward the center of the thick filament. This
process is referred to as crossbridge cycling.

The string of green circles

represents an actin filament.
There are binding sites in the
filament for the attachment of
myosin heads.
*In a relaxed muscle the
binding sites are covered by
tropomyosin. The
tropomyosin has molecules of
troponin attached to it.
*Calcium, shown in yellow, will
attach to troponin.
*Calcium will change the
position of the troponin,
tropomyosin complex. *The
troponin, tropomyosin complex
has now moved so that the
binding sites are longer covered by the troponin, tropomyosin complex.
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7) Sliding-Filament mechanism
Each cross bridge cycle consists of the following steps:
Binding (attachment) of
the cross bridge to active
sites of actin filaments.
Bending (movement) of
the cross bridge, produces
movement of the actin
filament inward.
Detachment of the cross
bridge from the thin
filament when the head of
Cross bridge automatically
breaks away from the
active sites and returns to
its normal position.
Then the cycle is repeated.

Mechanism of muscle
relaxation
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1- It is an active process. Ca++ is removed from the cytoplasm by Ca ++ pumps

located on the sarcoplasmic reticulum membrane.
2- When the intracellular Ca++ concentration in the myofibril cytoplasm, falls,
troponin return to its original state.
3- Tropomyosin moves back to cover the binding sites on actin.
4- Cross-bridge cycling stops, ending the contraction and allow muscle fiber
to relax.
5- Stop interaction between actin and myosin.

Skeletal Muscle relaxation is active due to two factors:

Energization of myosin head: binding of ATP to myosin head to bring it back to resting
state.
Active Ca reuptake by longitudinal tubules (by Ca2+ ATPase).
Muscle fibers of a dead body do not produce any ATP. So, after death:
Ca2+ is no longer pumped back into SR
ATP reserves needed to break down stable A-M complexes are depleted.
This results in stiffening of the dead body or rigor (firmness) mortis, which passes only
after the actin & myosin molecules in the muscle fibers decompose.
MYOSIN
Myosin I: present in sperms.
Myosin II: present in sarcomere (skeletal muscles).

Source of energy for muscular contraction

The immediate and direct energy source for muscle contractions is ATP.
Hydrolysis of ATP can proceed anaerobically.
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ATP ADP + Pi + energy.

Muscle has three pathways to generate ATP:

1- Creatine phosphate
- Creatine phosphate + ADP ATP + creatine.
- Frist few seconds of exercise.
- Needed for high intensity, short exercise as jumping. (for a 100-m sprint).

2- Anaerobic glycolysis
- Glucose + 2 ATP Anaerobically lactic acid + 4 ATP
- In absence of O2 , glucose is broken into lactic acid, with liberation of a
small quantity of ATP (2 ATP).
- frist few minutes of exercise.
- Continuation of anaerobic glycolysis may lead to excess lactic acid which
producing fatigue.
3- Oxidation phosphorylation
- Slow regeneration of ATP by energy derived from breakdown of glucose
in the presence of O2 .
- liberating a great amount of ATP (38 ATP).
- in prolonged exercise (Walking and swimming).
- It is a long term aerobic system.

During light exercise: lactate is broken down in the heart & liver whereby H+ ions are
used up.
During strenuous exercise: Anaerobic glycolysis must be continued along with aerobic
oxdn if it does not supply sufficient quantities of ATP.
For sustained exercise: Aerobic regeneration of ATP from glucose (about 32 ATP per
glucose residue) or fatty acids is required.

The Oxygen Debt Mechanism

Definition
the extra amount of oxygen that must be taken into the body during recovery
period after muscular exercise.
AIM
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to restore all the metabolic systems back to their full normal state.
1) A lactic oxygen debt
Replenish the ATP, O2 stores of the body and phospho-creatine stores.
2) lactic oxygen debt
To remove the excess lactic acid from muscles and all body fluids.

Muscle fatigue
Contractile activity in a skeletal muscle cannot be maintained at a give
level indentifintely. The tension in the muscle declines as fatigue sets in
following continuous stimulation of a muscle at high frequency.

Causes of fatigue according to its site

1) Muscular fatigue:
i) Accumulation of lactic acid, which may inhibits the key enzymes in
energy producing pathways or oxidation contraction coupling decreasing
Ca++ released from S.R.
ii) Depletion of energy reserves e.g. glycogen.
2) Neuromuscular fatigue:
It is due to depletion of acetylcholine at neuromuscular junction.
3) Central or psychological fatigue:
Occurs when the central nervous system (CNS) no longer adequately
activates the motor neurons supplying the working muscles.
The person slows down or stop exercising even through the muscles are still
able to perform may be due to decrease the individuals motivation to
continue the exercise.

During exercise: contraction of muscle squeezes the blood vessels.

Veins are squeezed impaired venous drainage so the metabolites & lactic acid
cannot escape.
[Exercise increases venous drainage for the veins in between the sk. Muscles.]
Since the arterial blood pressure is high, it can overcome the sk. Muscle pressure & flow
thru them.

Accumulation of metabolites (due to protein breakdown & lactate production) in the

sarcomere osmotic pressure extraction of water from ECF swelling blood
flow reflex tension pain.
Types of skeletal muscle fibers
Based on contraction time, skeletal muscles are classified into two types:

Feature Red (slow) muscles Pale (fast muscles)

Myoglobin High, so it is red ( for short- Less
content term O2 storage)
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Blood supply Rich Relatively Less

Mitochondria Rich Relatively Less
Sarcoplasmic Less extensive More extensive
retiuculum
Power of Less powerful More powerful
Contraction
Duration of Sustained contraction (longer Brief and rapid contractions.
contraction contraction time)
Fatigue Occurs slowly Occurs quickly
Energy Depends on cellular Depends on glycolysis
source respiration (aerobic) (anaerobic)
Have conc. of fat droplets Rich in glycogen.
(high-energy substrate
reserves)
Rich in oxidative enzymes
Examples Back muscles & Soleus (for upright position),
gastrocnemius Hand muscles & ocular
muscles

Each fiber type can also be converted to the other type.

If, prolonged activation of fast-twitch fibers leads to a chronic in
cytosolic Ca2+ conc. fast-twitch fibers will be converted to slow-twitch
fibers & vice versa.
Parvalbumin:
It is a protein occuring in the cytosol of fast-twitch muscle fibers.
Accelerates muscle relaxation after short contractions by binding
cytosolic Ca2+ in exchange for Mg2+.
Its binding affinity for Ca2+ is higher than that of troponin, but lower
than that of SRs Ca2+-ATPase.

Types Of Skeletal Muscle Contraction

Isometric contraction occurs when the muscle develops tension and exerts
force on an object, but does not shorten.
For example, supporting an object in a fixed position, such as carrying a
book. This type of contraction also occurs when attempting to move an object
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that is too heavy. In this case, the muscle may exert maximal force against the
object; however, because the object does not move, the length of the
contracting muscle does not change.

Isotonic contraction: occurs when the muscle shortens under a constant load.
For example, when an object is lifted, the muscle contracts and becomes
shorter although the weight of the object remains constant. In addition to
moving external objects, isotonic contractions are performed for movements
of the body, such as moving the legs when walking.

Isometric contraction Isotonic contraction

Shortening of central contractile
part = lengthening of peripheral
non-contractile parts
Total length is constant
E.g.: During upright position in
lower sk muscles to maintain the
body posture.
No external work done.
Shortening of central contractile part as
well as shortening & stretching of
peripheral ends.
Total tension is constant.
Could be associated with shortening or
lengthening (Lifting or placing down of
object with a constant tension (eg:
picking of precious objects very slowly).
External work done

Isotonic contraction Isometric contraction

N.B, Skeletal muscle formed of:

Central contractile part contains sk. Muscle fibers.
Peripheral non-contractile ends elastic non-contractile stretchable tissues
Contraction of sk muscle shortening of central contractile part (In the form
of shortening of sarcomeres.
Relation between Muscle Length and
Tension (Pre-loading)
Starling law:
Within limit, the strength of muscle contraction is directly proportionate to the
initial length of the muscle.
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Explanation:
When the muscle fiber contracts isometrically, the tension developed is
directly proportionate to the number of cross-linkages formed between actin
and myosin molecules.
(1) With very short initial length, the actin filaments will be crowded at the
center of sarcomere, which interferes with force generation.
(2) Increasing the length of muscle fiber will increase the length of
sarcomere, leading to better interaction between actin and myosin (with
more number of cross-linkages formation).
(3) L-max: it is the optimal sarcomere length (2 - 2.2 Um) at which the
active tension is maximal. At this length, maximal tension developed.
Above the optimal length , the actin filaments start to be pulled away from
the heads of myosin. Thus, muscle tension developed will be decreased.
(4) With further increase in muscle length (i.e. severe stretch), actin
filaments become completely pulled away from the myosin heads with
failure of contraction.

Simple muscle twitch

1- A single action potential in a muscle fiber produces a brief weak
contraction known as a twitch.
2- The time for action potential is about 1 to 2 msec and it is completed
before the start of muscle contraction.
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3- The time delay between stimulation

and onset of contraction is known as
the latent period during which
excitation contraction coupling occurs.
4- The time from the onset of
contraction until peak tension is
developed is the contraction time.
5- The time from peak tension until
relaxation complete is the relaxation
time.

Effect of Repetitive Stimulation on the skeletal muscle

The response of a skeletal muscle to multiple successive stimulation

depends on the frequency (rate) of stimulation as follows:
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1. If the frequency of stimulation is low, so that the

stimuli fall soon after the relaxation phases of the
preceding twitches, so separate twitches showing
Treppee (=staircase) phenomenon: is an increase in
the tension developed during each twitch until, after
several contractions, a uniform tension per
contraction is reached. Treppe is believed to be due
to the availability of Ca.

2. If the frequency of stimulation is increased so

that the stimuli fall during the relaxation phases of
the preceding twitches incomplete tetanus = clonus
is obtained. (= Contraction + incomplete
relaxation).

3. If the frequency of stimulation is further

increased so that the; stimuli fall at the contraction
phases of the preceding twitches complete tetanus is
obtained (= Continuous muscle contraction
without relaxation).
[It occurs at 20 Hz in slow-twitch muscles & at 60100 Hz in fast-twitch muscles].

N.B.
Tetanus: summation of contraction without relaxation due to marked in the
frequency of excitation.

Tetany: neuromuscular excitability due to hypocalcemia caused by

hypoparathyroidism etc

Treppe or Staircase Phenomenon

Gradual increasein force of contraction of muscle when it is stimulated repeatedly with a
maximal strength at a low frequency. It is different from summation & tetanus.
The action potential developed in muscle membrane is always of the
same magnitude (all-or-none). The activation of the contractile elements is
not. It is dependent on the amount of Ca++ released from the cisterns.
Successive stimuli produces action potentials which cause more and more
Ca++ to be released, thus increasing the contractile response.

Physical Fitness and Training

Physical exercise capacity can be measured by using ergometry.
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Ergometry: assesses the effects of exercise on physiological parameters such

as respiration rate, HR & plasma lactate conc.

Measured physical power (performance) is expressed in: watts (W) or

W/kg body weight (BW).

Short-term performance tests (1030 s): measure performance achieved

thru rapidly available energy reserves (CrP, glycogen).
Medium-term performance tests: measure performance fueled by
anaerobic glycolysis.
Longer term aerobic exercise performance: measure performance fueled
thru oxdn of glucose & FFA by measuring V.O2 max.

In strenuous exercise (2/3 the max. physical capacity or more), aerobic

mechanisms do not produce enough energy, so anaerobic metabolism must
continue as a parallel energy source lactacidosis.

Physical training
Raises & maintains the physical exercise capacity.

There are 3 types of physical training strategies:

Endurance training Strength training
Improves submaximal long-term Improves max. short-term
performance performance level
e.g., running a marathon e.g., in weight lifting
To the oxidative capacity of To muscle mass by the size of the
slow-twitch motor units. muscle fibers (hypertrophy) & to the
glycolytic capacity of type F motor
units.

Trained athletes: can therefore achieve larger increases in CO & ventilation.

In individuals practicing endurance training, exercise-related rise in the
lactate conc. is also lower & occurs later than in untrained.

Excessive physical exercise causes muscle soreness & stiffness (not due
to lactic acid accumulation), but sarcomere microtrauma muscle swelling
& pain.
Physical Work

There are three types of muscle work:

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* +ve dynamic work: requires muscles involved to alternately contract &

relax (e.g., going uphill).

* -ve dynamic work: requires muscles involved to alternately extend while

braking (braking work) & contract without a load (e.g., going downhill).

* Static postural work: requires continuous contraction (e.g., standing

upright).

Many activities involve a combination of 2 or 3 types of muscle work.

Outwardly directed mechanical work is produced in dynamic muscle activity,
but not in purely postural work.

Purely postural work

Force x distance = 0
Chemical energy is still consumed & completely transformed into a form of
heat called maintenance heat (= muscle force x duration of postural work).
in blood flow is prevented as the continuously contracted muscle squeezes
its own vessels. The muscle then fatigues faster than in rhythmic dynamic
work.

Effects of Exercise on Skeletal Muscles

Our body is able to adapt to almost any kind of stimulus or stress.

Muscle Structure
The look of your muscle changes depending on the type of exercise
you are doing. Endurance training such as jogging is exercise with a small
force but a more frequent activation of your skeletal muscle cells. This type
of training leads to an increase in the metabolic rate of your muscle and less
in the size and shape. Resistance training such as weightlifting is a high-force
exercise with less frequent muscle activation. Long-term effects of resistance
training lead to hypertrophy of the skeletal muscle, an increase in cross-
sectional size.

Size and Nerve Function

When you lift weights or engage in other muscle-building activities,
you increase the amount of myosin and actin inside the fibers of each affected
muscle; in turn, this increase causes your muscles to gain size or mass. This
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process, which typically takes place gradually over a period of weeks or

months, is known as hypertrophy. Regular participation in exercise also
increases the efficiency of the nerve signals that pass between your muscles
and your brain and allow you to consciously and accurately control your
movements.

Muscle Fiber Activity

By engaging in certain types of activities, you can control exercise's
effects on your fast- and slow-twitch muscle fibers. For instance, regular
performance of activities that require aerobic endurance will lead to an
increase in the size of your fast-twitch muscle fibers. If you perform
moderate types of muscle-building, you will increase the size of both your
fast- and slow-twitch muscles. However, if you engage in intense muscle-
building activities, you will only increase the size of your fast-twitch
muscles. Despite common conceptions, engaging in certain forms of exercise
will not allow you to convert fast-twitch fibers into slow-twitch fibers or vice
versa.

Anti-Aging Effects
Long-term regular performance of muscle-building exercises can help
partially offset age-related loss of skeletal muscle in older individuals.

During physical exercise:

Feature Resting value During exercise (max. value)
Ventilation (V.E) 7.5 L/min 90 to 120 L/min
Respiratory rate 4060 min1max
Tidal volume 2L
O2 consumption (V.O2) 0.3 L/min 3 L/min (due to O2 consumption
in tissues)
Pulmonary transit time 0.75s 0.25s (Time less than 0.25s, no
(Minimal time sufficient proper gas exchange)
for gas exchange)
O2 consumption: calculated as the arteriovenous difference in O2 conc. =
avDO2 (L/L blood) x Blood flow (L/min)
Maximum O2 consumption (V.O2 max) is defined as:
V. O2 max = HRmax SVmax avDO2max
Ideal measure of physical exercise capacity: V.O2 max per body weight
Functional differences between cardiac
muscle & skeletal muscle:
Skeletal muscle Cardiac muscle
More extensible Less extensible, so the passive extension force at
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Excitable Tissues

rest is greater.
Functions in the Operates in the ascending limb (below Lmax) of its
plateau region of its lengthforce curve without a plateau.
lengthforce curve. So, the ventricle responds to diastolic filling loads
by its force development (FrankStarling
mechanism).
Extension also affects troponins sensitivity to
Ca2+ steeper curve
APs are of shorter APs are of much longer duration bcoz gK &
duration. gCa temporarily after rapid inactivation of Na+
Uses IC Ca2+, so no channels slow influx of EC Ca2+ plateau
plateau phase. phase of AP.
As a result, the refractory period does not end until
a contraction has almost subsided. So, tetanus
cannot be evoked in cardiac muscle.
Contains motor units. Has no motor units. The stimulus spreads across
all myocardial fibers of the atria & ventricles all-
or-none contraction of both atria & thereafter,
both ventricles.
No change Duration of AP can change the force of contraction
(which is controlled by the variable influx of Ca2+
into the cell).

Smooth Muscle
1) They contain myosin filament and actin filament, no troponin in thin
filaments. (instead calmodulin present).
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Excitable Tissues

2) The contractile proteins are not regularly arranged with no apparent

sarcomeres or striations.
3) No apparent Z lines.
4) Smooth muscle cells do not have T tubule and SR poorly developed.
5) It contains few mitochondria and it depends to a large extent on
glycolysis for their metabolic needs.
6) Smooth muscles are un-striated, involuntary slow contracting supplied by
autonomic nerves.
7) It can show maintained prolonged contractions without fatigue and with
little energy consumption.
Excitation-Contraction Coupling in Smooth muscle
1-Calcium influx into the cytoplasm by
opening the voltage-sensitive and ligand
gated Ca++ channels. (90% of Ca from
ECF and 10% from SR).
2-After the increase in cytosolic Ca ++,
Ca++ binds to cytoplasmic protein
Calmodulin.
3-Ca++ Calmodulin complex activate
myosin light chain kinase (MLCK).
4-The activated (MLCK) uses ATP to phosphorylate the myosin cross-
bridges allowing myosin ATPase to be activated.
5-The phosphorylated myosin cross bridges bind to actin and undergoes
Walk-along cross bridge cycling.

Mechanism of relaxation: active process

+
1. To cause relaxation of the smooth muscle, it is necessary to remove the Ca
+
ions.
++ ++
2. This occurs by Ca pump, which pumps Ca out of the smooth muscle
fiber and back into the ECF or pumps Ca++ intro S.R.
3. Also to relax the contracted smooth muscle the phosphorylated myosin
must be dephosphorylated by phosphatase enzyme.

Thermal changes
The liberation of energy during muscle contraction is associated with the
production of heat.
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Excitable Tissues

(a) Resting heat - heat given off by the inactive muscle, it is due to the
metabolic process of the resting muscle.

(b) Initial heat heat liberated during the active phase of contraction. It is
made-up of the following:
(i) Activation heat - heat production when the muscle
contracts as in a muscle twitch. (in titanic contraction it is called maintenance
heat)
(ii) Hat of shortening - liberated when the muscle shortens. It is dependent
upon the degree of shortening.

(c) Recovery heat or delayed heat

Additional amount of hat liberated as a consequence of metabolic processes
which returns the muscles into its original precontraction
physical condition.

(i) Heat of relaxation - liberated when isotonic muscle contraction is over and
relaxation occurs. The heat is produced as a result of the structuring by the
load