Mechanics of Muscles

170301 Biomechanics

Prof. B. H. Parmar
bhparmar@gmail.com
Eccentric and Concentric Contractions
• Concentric: Energy • Eccentric: Energy
generating, Positive Work, absorbing, Negative work,
accelerating braking
Increase joint angular Increase joint angular
velocity and increase the velocity and reduce the
total energy of the total energy of the
system system
Positive and Negative Joint Work
• direction of joint work
depends on polarity of
joint torque and
direction of joint
Rotation
• movements 2, 3, 4, and 5
involve positive work or
concentric Contractions
• movements 1, 6, and 7
involve negative work
or eccentric
contractions
 Types of Muscles
• There are three types of muscles:
–skeletal,
–smooth, and
–cardiac.
• Smooth muscles line the internal
organs, and cardiac muscles form the
heart.
 Skeletal Muscles

• Skeletal muscles is attached,

via aponeuroses / tendons, to
at least two bones causing /
controlling the relative
movement of one bone with
respect to the other.
 Skeletal Muscles
• When its fibers contract under the
stimulation of a nerve, the muscle
exerts a pulling effect on the bones to
which it is attached.
• Contraction is a unique ability of the
muscle tissue, which is defined as the
development of tension in the muscle.
 Skeletal Muscle
• Muscle contraction can occur as a
result of
– muscle shortening (concentric
contraction) or
– muscle lengthening (eccentric
contraction), or
– occur without any change in the muscle
length (static or isometric contraction).
 Skeletal Muscle
Actin

Myocin
• The skeletal muscle is
composed of muscle fibers and
myofibrils.
• Myofibrils in turn are made of
contractile elements: actin and
myosin proteins.
• Actin and myosin appear in
bands or filaments. Cross-bridge
 Skeletal Muscle
Actin

Myocin
• Several relatively thick
myosin filaments interact
across cross-bridges with
relatively thin actin filaments
to form the basic structure of
the contractile element of the
muscle, called the sarcomere
Cross-bridge
 Skeletal Muscle
Actin

Myocin
• Many sarcomere
elements connected in
a series arrangement
form the contractile
element (motor unit)
of the muscle.
Cross-bridge
 Skeletal Muscle
• It is within the sarcomere that the Actin

Myocin
muscle force is generated, and where
muscle shortening & lengthening
takes place.
• The active contractile elements of the
muscle are contained within a fibrous
passive connective tissue, called
fascia.
• Fascia encloses the muscles, separates
Cross-bridge
them into layers, and connects them
to tendons.
 Muscle Force Results
from Interaction between
Contractile Proteins
• sarcomere: the smallest
anatomical unit that contracts like
a muscle
• sliding filament model proposes
that muscle force arises from
cyclic binding between thick and
thin filaments of the sarcomere
• thin filaments contain actin,
troponin C, and tropomyosin
• thick filaments contain myosin
• in the absence of calcium,
tropomyosin prevents myosin
from attaching to actin
 Calcium is Needed for Muscle
Contraction
• at the onset of an action potential,
the sarcoplasmic reticulum (SR; a
membrane that surrounds the
myofibrils) releases calcium
• calcium binds to troponin, causing
a conformal change in
tropomyosin which reveals myosin
binding sites on the actin
• simultaneously, adenosine
triphosphate (ATP) is hydrolyzed
by ATPase in the myosin head,
providing the energy for cross-
bridge Attachment
• the SR re-sequesters calcium at
the end of the action potential,
thereby inducing muscle
relaxation
 Factors Affecting Force and Torque
Developed by Muscle
• The force and torque developed by a muscle is
dependent on many factors, including
– Number of motor units within the muscle,
– Number of motor units recruited,
– Manner in which the muscle changes its
length,
– Velocity of muscle contraction, and
– Length of the lever arm of the muscle force.
 Factor Affecting Muscle Force
development (a partial list)
• Muscle Fibre Type
• Number of Activated Motor Neurons,
Frequency of Discharge
• Muscle Length
• Velocity of Shortening / lengthening
• Muscle Geometry (Physiological cross-
sectional area (PCAS), angle of pennation)
 Muscle-nerve Interaction
• a motor nerve enters
muscle and splits into
numerous axons; each
axon contacts 10- 2000
muscle fibres
• each muscle fibre is
innervated by only one
motor nerve axon, and
contracts in response to
an action potential in that
axon
• motor unit: a single motor
nerve axon and all the
muscle fibres it contacts
 Stimulation Frequency Affects
Muscle Force: Twitch and Tetanus
• muscle force can be modulated
by varying: (1) the number of
recruited motor neurons, and
(2) the frequency of discharge
(i.e., stimulation rate) in motor
neurons
• a single action potential (S1)
produces a twitch contraction, a
quick rise and slow fall in force
• a tetanus occurs when a new
action potential (S2) arrives
before the previous twitch has
dissipated, and there is force
summation
• at stimulation frequencies >30/s,
there are no twitch transients
(fused tetanus)
 Three Types of Muscle Fibres and
Motor Units, Defined by Contraction
Speed, Peak Force, Fatigue Resistance
Tetanus 40 s-1 Twitch Fatigue test Fibre type Motor unit
type
SO (Slow, S (Slow)
Oxidative)

FOG (Fast, FR (Fast,

Oxidative, Fatigue-
Glycolytic) resistant)

FG (Fast, FF (Fast,
Glycolytic) Fatiguable)
 Size Principle
• When a stimulus is applied to
the ventral aspect of the spinal
cord, the smallest and most
excitable motor units are
activated first. These tend to
be slow (S) motor units which
innervate slow oxidative (SO)
muscle fibres. Larger FR and
FF motor units that innervate
FOG and FG fibres are
recruited only at high levels of
force.
• Sequence is reversed when
force level falls, with largest
motor units dropping out first.
 Active Force Development in the
Sarcomere Depends on Actin-Myosin
Overlap
• (A): no overlap between actin
and myosin, zero developed
Tension
• between (A) and (B): tension
increases linearly as overlap
Increases
• between (B) and (C): maximum
overlap & maximum tension
• left of (C): interference between
actin filaments reduces ability of
crossbridges to develop tension
• left of (D): myosin filaments
collide with Z-lines and fold,
and force declines rapidly
 Factors Affecting Force and Torque
Developed by Muscle
• For muscles, two different forces can be
distinguished.
– Active tension - Force produced by the contractile elements of the
muscle (result of voluntary muscle contraction)
– Passive tension - Force developed within the connective muscle
tissue (when the muscle length surpasses its resting length)

• The net tensile force in a muscle is

dependent on the force–length
characteristics of both the active and
passive components of the muscle.
 Factors Affecting Force and Torque
Developed by Muscle
• The number of cross-
bridges between the
filaments is
maximum, and
therefore, the active
tension (Ta) is
maximum at the
resting length (lo) of
the muscle.
 Factors Affecting Force and Torque
Developed by Muscle
• As the muscle lengthens, the
filaments are pulled apart,
the number of cross-bridges
is reduced and the active
tension is decreased.
• At full length, there are no
cross-bridges and the active
tension reduces to zero.
• As the muscle shortens, the
cross-bridges overlap and the
active tension is again
reduced.
 Factors Affecting Force and Torque
Developed by Muscle
• When the muscle is at its
resting length or less, the
passive (connective)
component of the muscle is in
a loose state with no tension.
• As the muscle lengthens, a
passive tensile force (Tp)
builds up in the connective
tissues.
 Factors Affecting Force and Torque
Developed by Muscle

• Passive tensile force

increases at an increasing
rate as the length of the
muscle increases.
• The total muscle force (Tt)
transmitted via tendons is
the sum of the forces in the
active and passive The force–length
elements of the muscle. characteristic of passive
component resembles that of
a nonlinear spring.
 Factors Affecting Force and Torque
Developed by Muscle
• Tension–length diagram is not unique
but dependent on the number of motor
units recruited.
• The magnitude of active component of
the muscle force can vary depending
on how the muscle is excited, and
usually expressed as the percentage of
the maximum voluntary contraction.
 Muscle Length Affects Force
Developed in whole Muscle
• the tension developed in a
whole muscle is the sum of
active force due to muscle
contraction and passive force
due the passive stiffness of
tendon and muscle
• the passive force is negligible
for lengths less that the normal
resting length (l0)
• the active force follows the
tension-length behaviour of the
sarcomere, and scales with
muscle activation
 Muscle Velocity Affects Force
Development in Whole Muscle
• force (T) is greater during
lengthening than
shortening Contractions
• the greater the shortening
velocity (v), the smaller the
force (explains why we
cannot lift heavy objects
quickly)
• in the shortening regime,
mechanical power output is
maximum when T and v
are around one-third their
maximum values
 Greatest Force Developed when
Lengthening near Resting Length
 Muscle force Transmission
• The force generated by a contracting
muscle is usually transmitted to a bone
through a tendon.
• There is a functional reason for
tendons to make the transfer of forces
from muscles to bones.
 Muscle force Transmission
• As compared to tendons, muscle have lower
tensile strengths.
• The relatively low ultimate strength
requires muscles to have relatively large
cross-sectional areas in order to transmit
sufficiently high forces without tearing.
• Tendons are better designed to perform this
function.
 Muscle Force-Velocity behaviour is
Described by the Hill Equation

• (T+a)(v+b)=(T0+a)b
– T0 - Isometric tension
– Vmax – Maximum Velocity
= bT0/a
• Instantaneous Power
P=T∙v
 Hill’s Active State Model of Muscle
Contraction
Assumptions
– for a given length,
muscle always
develops the same
peak force T0(x1,t);
– if the muscle is
shortening, some
force is dissipated in
overcoming inherent
viscous resistance
 Hill’s Active State Model of Muscle
Contraction

• B: muscle damping constant,

– must be a nonlinear function of shortening velocity and
temperature
• KSE: stiffness of the series elastic component;
– represents force-deflection properties of tendon
• KPE: stiffness of the parallel elastic component;
– represents force-deflection properties of sarcolemma, epimysium,
perimysium, and endomysium
 Quick-release Experiments for
Determining the Hill Model
Parameters
Steps:-

1. Hold muscle length fixed with

the catch
2. Stimulate muscle to produce
peak (isometric) force T0
3. At the instant of release,
muscle force is reduced to a
value T (where T < T0) that
depends on weight in pan
Quick-release Experiments (Cont…)
1. there is an instant change
(Δx2) in the length of KSE
following release
2. Followed by a more gradual
change (Δx1) in the length of
the muscle
3. As T increases, there is a
decrease in v (slope of dashed
line), reflecting that muscle
cannot shorten quickly under
high loads
4. Combinations of T and v
reflect the force-velocity
properties of a given muscle
 Shortcoming of the Active State
Model
• Predicts a negative T0 during the
end phase of the twitch (marked
by asterisk in diagram at right)
• The liquid in muscle (water)
does not have the required
nonlinear damping
characteristics
• Cannot accurately predict
muscle force during lengthening
(the slope of the T-v curve is
about 6-fold greater for
lengthening than shortening,
and muscle length increases
rapidly when the load exceeds
1.8T0)
Huxley’s Sliding Filament Model
• Force development is due to
stretch of elastic myosin
crossbridges, which can form
bonds with actin for x<h
• Bonds can be maintained for
x>h (tensile) and x<0
(compressive)
• For (0<x<h), rate of
attachment (f) exceeds rate of
detachment (g)
• Rate of detachment (g) is
slower during lengthening
than shortening, thus
accounting for greater force
under eccentric conditions