GRADATION OF

MUSCULAR
ACTIVITY
Learning objectives

• Motor unit
• Innervation ratio
• Motor unit types – I,II
• Elastic elements of skeleton muscle & force generation
• Active tension & passive tension
• Isometric length tension relationship
• Isotonic force velocity relationship
• Summation – Multimotor unit summation
- Frequency summation
 Motor unit

Definition

A motor unit consists of a single motor neuron,

its axonal branches and all the muscle fibers
supplied by them
Structure of a motor unit
Size of a Motor Unit

• The cell bodies of motor neurons are present in the

anterior horn of the spinal cord.
1. The size of a motor unit can be large or small
depending on the number of muscle fibers innervated by
it.
2. In the intrinsic muscles of the hand, one motor neuron
innervates less than ten muscle fibers so, stimulation of a
motor unit produces a small rise in tension.
3. The back muscles have hundreds or even thousands of
muscle fibers per motor unit, where activation of a single
motor unit causes a large increase in tension.
Innervation ratio

• The number of muscle fibres innervated by a single motor neuron is

called its innervation ratio
• A single motor neuron branches out and innervates many
muscle fibers.
• When a motor neuron discharges, contraction is produced in
all the muscle fibers supplied by it
Recruitment of Motor Units

• At the resting state of the muscle, hardly any motor unit is

activated.
• When a single motor neuron discharges, there is some rise in
tension in the muscles of the concerned limb.
• If a greater total tension has to be attained, additional motor
neurons fire and there is increase in the number of active
motor units.
• The process of activating more number of motor units is
called recruitment of motor units
• Application :
Asynchronous discharge of motor units prevents fatigue:
The motor units discharge asynchronously.
When some of the motor units are active, other units are
silent.
Afterward, the active units go to rest and the silent units
become active.
At any instant of time, the inputs from different units
are summated and this results in a smooth contraction of
the muscle.
This also helps the muscle to work for longer time
without being fatigued.
 Types of skeletal muscle fibres
• Muscle fibers can be classified as fast twitch or slow twitch.
• The type of fiber is determined by its innervation.
• All fibers in a motor unit are innervated by a single α motor neuron.
• All fibers within a motor unit are of the same type.
 Types of skeletal muscle fibres

• Slow-twitch motor units tend to be small (100 to 500 muscle

fibers) and are innervated by an α motor neuron that is easily
excited.
• Fast-twitch motor units tend to be large (containing 1000 to 2000
muscle fibers) and are innervated by α motor neurons that are
more difficult to excite.
Types of skeletal muscle fibres

• Slow-twitch motor units tend to be recruited first.

• As more and more force is needed, fast-twitch motor units are
recruited.
• The advantage of such a recruitment strategy is that the first muscle
fibers recruited are those that have high resistance to fatigue.
• Moreover, the small size of slow-twitch motor units allows fine motor
control at low levels of force.
 Types of motor units

• Type I- Red or slow Motor units

• Type II- White or fast Motor units

Characteristics Type I Type II
Muscle fibre type Slow, red, involved in tonic activity Fast, white, involved in phasic
activity

Metabolism Aerobic, low glycolytic, high Anaerobic, high glycolytic, low

oxidative capacity oxidative capacity

Number of mitochondria High Low

Glycogen content Low High

Capillary density High Low

Blood supply High Normal

Myoglobin content High Low

Enzymes

Phosphorylase activity Low High

Myosin ATPase activity High Low

NADH Dehydrogenase Low High

Axon diameter Small Large

Conduction velocity Slow Fast

Characteristics Type I Type II

Twitch duration of the muscle Long Brief

Tetanic tension Small Large

Types of movements Tonic contractions Phasic contractions

Eg: Posture Eg: Skilled movements
First to be recruited during Remain inactive during
muscle contraction contraction
Recruited only when powerful
contraction is required

Fatigability Fatigue resistant Easily fatigable

Further types - II a- fast fatigue resistant,

glycolytic
II b- fast, fatigable, glycolytic
Properties of motor units
 Elastic elements and force generation

• Parallel elastic element-

Sarcolemma
• It is non contractile.
• Lies in parallel to muscle fibres.
• Provides resistance when stretched.

• Series elastic element- Tendons,

hinge regions of myosin.
• Stores energy when stretched
 Elastic elements and force
generation
• If a muscle is rapidly stretched during the early part of a
contraction, it is capable of developing a considerably greater
tension than that recorded under normal condition(without
stretching).

• This reveals the presence of an elastic element coupled in series

with the contractile element.
Experimental evidences

• Hill- Quick stretch experiment- much of the elasticity resided in the

tendons and in the connective tissues in series with contractile
elements.

• H.F.Huxley- Site of series elasticity as well as the site of tension

generation are both in the sarcomere.
• Cross bridges, hinge regions of myosin are the sites of tension
generation in the muscle. They are independent force generators.
• Degree of overlap between thick and thin filaments is directly
proportional to the level of tension generated in a twitch.
 Active, passive and total
tension
• Tension in a stimulated muscle- Total tension
• Tension in an unstimulated but stretched muscle- Passive tension
• Difference between total and passive tension- Active tension
Isometric length
tension
relationship
According to
starlings law –
force of
contraction is a
function of the
initial length of
muscle fibre
Isotonic force velocity
relationship
Summation
• Summation- adding together of individual twitch contractions
to increase the intensity of overall muscle contraction.
It occurs in two ways:
• (1) by increasing the number of motor units contracting
simultaneously- multiple fiber summation
• (2) by increasing the frequency of contraction, which is
called frequency summation and can lead to tetanization.
 Multiple Fiber Summation
• When the central nervous system sends a weak signal to contract a
muscle, the smaller motor units of the muscle are stimulated in
preference to the larger motor units.
• As the strength of the signal increases, larger and larger motor units
begin to be excited.
• The largest motor units often having as much as 50 times the
contractile force of the smallest units. - the size principle.
• Importance - it allows the gradations of muscle force during weak
contraction to occur in small steps. The steps become progressively
greater when large amounts of force are required.
Size Principle

• Henneman's size principle states that under load, motor units are
recruited from smallest to largest.
• In practice, this means that slow-twitch, low-force, fatigue-resistant
muscle fibers are activated before fast-twitch, high-force, less fatigue-
resistant muscle fibers.
• The advantage of such a recruitment strategy is that the first muscle
fibers recruited are those that have high resistance to fatigue.
• Cause of size principle - the smaller motor units are driven by
small motor nerve fibers, and the small motor neurons in the
spinal cord are more excitable than the larger ones, so they are
excited first.
• Different motor units are driven asynchronously by the spinal
cord, so contraction alternates among motor units one after the
other, thus providing smooth contraction even at low
frequencies of nerve signals.
Frequency Summation and
Tetanization
• At low frequency of stimulation individual twitch contractions occur
one after another.
• As the frequency increases, each new contraction occurs before the
preceding one is over.
• As a result, the second contraction is added partially to the first, so
the total strength of contraction rises progressively with increasing
frequency.
• When the frequency reaches a critical level, the successive
contractions become so rapid that they fuse together and the whole
muscle contraction appears to be completely smooth and continuous-
tetanization.
The Staircase Effect (Treppe)

• When a muscle begins to contract after a long period of rest, its initial
strength of contraction may be as little as one-half its strength 10 to
50 muscle twitches later.

• The strength of contraction increases to a plateau, a phenomenon

called the staircase effect, or treppe.
Frequency Summation and
Tetanization
Thank u