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Physics Of Muscle

Amar Alansi
M.Sc BioPhysics
Microscopic Structure of A Myofibril
Each myofiberil consists of a number of two alternating
bands which are also called the sections, segments or
These bands are formed by muscle proteins
The two bands are
1- Light band or I band
2- Dark band or A band
Light band : its called I (Isotropic) band because it is
isotropic to polarized light. When polarized light is passed
through the muscle fiber at this area, light rays are
refracted at the same angle
Dark band: its called A (Anisotropic) band because it is
anisotropic to polarized light. When polarized light is
passed through the muscle fiber at this area, the light rays
are refracted at different dirctions. (An= not, iso= it,
trops= turning). Its called Q disk (Querscheibe= cross disk)
I band and A band are placed side by side
I band is divided into two portions, by means of a narrow and dark line called
Z line ( in German, zwichenscheibe = between disks)
The Z line is formed by a protein disk, which does not permit passage of lighe.
Sarcomere is portion of myofibril in between two Z lines
Its defined as the structural and functional unit of a
skeletal muscle, also called the basic contractile unit of the
Each sarcomere extends between two Z lines of myofibril,
its arranged in series
When the muscle is relaxed state, the average length of
each sarcomere is 2-3
In the middle of A band there is a light area called H zone
In the middle of H zone lies the middle part of myosin
filament this is called M line and M line is formed by myosin
binding proteins
Electron Microscopic study of
Sarcomere consists of many thread like structures called myofilaments
Myofilaments are of two types
1- Actin filaments
2- Myosin filaments
Actin filaments : its thin filaments with a diameter of 20 A and a length of 1
, these filaments extend from either side of the Z lines, run across I band
and enter into A band up to H zone
Myosin filaments: its thick filaments with a diameter of 115A and a length of
1.5 , these filaments are situated in A band

Myosin filaments are formed by myosin molecules

Actin filaments are formed by three types of proteins (Actin, Tropomyosin ,
Each myosin molecule has two poetions
1- Tail portion
2- Head portion
Each actin molecule is called F-actin
Each F-actin molecule has an active site to which the myosin haed is attached
The Tropomyosin molecules cover all the active sites of F-actin molecules
Troponin is formed by three subunits
1- Troponin I: wich is attached to F-actin
2- Troponin T : which is attached to tropomyosin
3- Troponin C: which is attached to clcium ions
Sliding Mechanism (ratchet)

During the contraction of the muscle:

the actin filaments glide/slide down between the myosin filaments
towards the center of H zone and approach the corresponding
actin filaments from the next Z line
The Z lines approach the ends of myosin filaments, so that the H
zone and I band are shortened during contraction of the muscle
During the relaxtion of the muscle:
the actin filaments and Z lines come back to the original position
Muscle Force
If the average diameter of the threads is D, the number of N per square centimeter of muscle is

The maximum pulling force Ff produced by the surface tension on each fiber is

The total maximum force Fm due to all the fibers in a 1cm2 area of muscle is

The average diameter D of the muscle fibers is about 100A (10-6cm). the maximum contracting
force that can be produced by surface tension per square centimeter of muscle area is

A surface tension of 1.75 dyn/cm can account for the 7.106dyn/cm2

The muscle produce about 30-100 N/cm2, speed of contraction is 0.5-1.6 m/sec, Fc31012 N
per cross bridge H.W?
Muscle Efficiency
ATP + H2O ADP + Pi + Q
Heat of reaction Q = h 6 1020 J/reaction (namely about 38 kJ/mol)
The cross-bridge head pushes the actin section by a distance of x 10 nm
This represents the work W = Fc x = 3 1012 N 108m = 3 1020 J
The difference QW 6 1020 J 3 1020 J = 3 1020 J must be carried off as heat
The efficiency of generating mechanical energy is ATP = W / Q 50%
The overall efficiency of the muscle one must include the metabolic process where glucose and oxygen
burned in order to convert ADP and Pi into the muscle fuel ATP. If this process has an energy efficiency
of about met=50%, then the overall efficiency of generating work from glucose is
=ATP met 0.5 0.5 = 25%.
The force generating muscle cycles may be compared to Carnot cycles of thermodynamic engines,
which have typical efficiencies of 20 to 30%.
If a muscle generates a force of Fc = 3 1012 N per cross bridge, and the muscle produces the stress
force 2 105 N/m2
there must be n = (2 105N/m2)/(3 1012N)=6.7 1016 cross bridges/m2.
Strength Duration Curve
Its also called Excitability curve
Its the graph that demonstrates the exact relationship between
the strength and the duration of a stimulus
Rheobase: its the minimum strength (voltage) of stimulus, which
can excite the tissue. The voltage below this can not excite the
Utilization time: its the minimum time required got rheobasic
strength of stimulus (threshold strength) to excite the tissue
Chronaxie: its the minimum time required for a stimulus with
double the rheobasic strength (voltage) to excite the tissue
Longer the chronaxie, lesser is the excitability
In human 0.08-0.32 msec, in frog 3 msec
Ten times more in skeletal muscles of infants than in that of adults
Types of contraction

Its classified into two types based on change in the length of muscle fibers or
tension of the muscle
Isotonic Contraction: its the type of contraction in which the tension remains the
same and the length of the muscle fiber is altered (ios= same, toinc= tension)
Ex. Simple flexion of arm, where shortening of muscle fiber occurs but the tension
does not change
Isometric Contraction: its the type of contraction in which the length of muscle
fibers remains the same and the tension is increased
Ex. Pulling any heavy object when muscles becomes stiff and strained with
increased tension but the length does not change
Simple Muscle Contraction
Its called twitch or curve
When the stimulus with threshold strength is applied, the
muscle contracts and then relaxes
Simple contraction of the muscle is called simple muscle twitch
and the graphical recording of this is called simple muscle curve
1- Point of stimulus (PS): the time when the stimulus is applied
2- Point of contraction (PC): the time when muscle begins to
3- Point of maximum contraction (PMC): the point up to which the
muscle contracts. It also indicates the beginning of relaxation of
the muscle
4- Point of maximum relaxation (PMR): the point when muscle
relaxes completely
1- Latent period: its the time interval between the point of
stimulus and point of contraction> the muscle does not
show any mechanical activity during this period
2- Contraction period: its the interval between point of
contraction and point of maximum contraction. The
muscle contracts during this period
3- Relaxation period: its the interval between point of
maximum contraction and point of maximum
relaxation. The muscle relaxes during this period
When the stimulus with threshold strength is applied,
the muscle contracts and then relaxes
Latent period : 0.01 sec
Contraction period : 0.04 sec
Relaxation period : 0.05 sec
Total twitch period : 0.10 sec
Contraction period is always shorter than relaxation
period. Its because the contraction is an active
process and relaxation is a passive process
Frank Starling Law
Work done in free loaded condition is more than in after loaded condition, this proves Frank
Starling law, i.e, the force of contraction is directly proportional to the initial length of muscle
fibers within physiological limits
Work done by the muscle
Work done = W h
W is weight lifted by the muscle
h is height up to which the weight is lifted
After load: its load that acts on the muscle after the beginning of muscular contraction. Example of
after load is lifting any object from the ground. The load acts on muscles of arm only after lifting
the objet off the ground, i.e. only after beginning of the muscular contraction
Free load: its the load in which acts on the muscle freely, even before the onset of contraction of
the muscle. Its otherwise called fore load . Example of free load is filling water from a tap by
holding the bucket in hand
Length Tension Relationship
Tension or force developed in the muscle during resting
condition and during contraction varies with the length of the
Tension developed in the muscle during resting condition is
known as passive tension. While tension developed in the
muscle during isometric contraction is called total tension
Active Tension: its the difference between the passive tension
and total tension at a particular length of the muscle. its
considered as the real tension that is generated in the muscle
during contractile process. It can be determined by the length
tension curve
Resting length: its the length of the muscle at which the active
tension is maximum. Active tension is proportional to the length
of the muscle up to resting length. Beyond resting length, the
active tension decreases
A micrometer is used to set length of the muscle fibers.
Force transducer is connected to a polygraph.
Polygraph is used to measure the tension developed by the muscle
during isometric contraction
Changes during Contraction
Optimal Length (l0)
Hill's muscle model
The three-element Hill muscle model is a representation of the muscle
mechanical response.
The model is constituted by a contractile element (CE) and two non-linear
spring elements, one in series (SE) and another in parallel (PE).
The active force of the contractile element comes from the force generated by
the actin and myosin cross-bridges at the sarcomere level.
The connective tissues that surround the contractile element influences the
muscle's force-length curve.
The parallel element represents the passive force of these connective tissues
and has a soft tissue mechanical behavior. The parallel element is responsible
for the muscle passive behavior when it is stretched, even when the contractile
element is not activated.
The series element represents the tendon and the intrinsic elasticity of the
myofilaments. It also has a soft tissue response and provides energy storing
Hill force velocity curve
The maximum tension Tmax that can be developed occurs for v=0 (isometric
T decreases and becomes zero at vmax
Hill force velocity
( T + a ) ( v+ b ) = ( Tmax + a ) b
Vmax= b Tmax / a
v' = ( 1 - T' )/( 1 + T' / k ) H.W?
Where v = v / vmax
T = T / Tmax
k = a / Tmax = b / vmax
Typically k values is 0.15 < k < 0.25
The power needed to move an object is F v, then the power generated by the
muscle is
P = T v = v ( b Tmax a v )/( v + b)

Cycle Time
There is a cycle time, cycle for a given cross bridge, during which time the myosin is
attached to the actin for a time on and is detached and inactive for a time off
cycle = on + off

During the on-time the cross bridge moves the attached

phase via hydrolysis by the power stroke distance or working
distance , which is about 5.3nm in vitro for skeletal muscle
myosin II.
The duty ratio rduty is the fraction of time that the cross bridge
is attached:
rduty = on /( on + off ) = on / cycle

For myosin on is 103 s and rduty is 0.02, so a given myosin

molecule is in contact with an actin filament for a relatively
small fraction of the time.

The minimum number of heads Nmin required for continuous movement

Nmin = 1 / rduty
With Nmin 1/0.02 = 50, only about six of the myosin heads of the roughly 300 myosin cross
bridges in a half a filament are attached and exerting a force at any given time.
During a complete cross bridge cycle, which is the time it takes to complete one ATP-
hydrolysis cycle, the motor moves a distance , where
= / rduty
Typically 5nm 50 = 250nm (for low loads).

The cycle time is related to the rate constant for the

hydrolysis of an ATP molecule kATPase
cycle = 1 / kATPase
Using kATPase 20s-1, we see that cycle 0.05 s

The relative speed of the sliding actin and myosin filaments is given by the total displacement
during the cycle time v, which is
v = kATPase = / cycle = / on
So we see that v = 5nm/103s = 5m/s.

In addition to and , there is a third characteristic distance, d, which is the path distance or
step size between consecutive binding sites, and is 36nm for this cross bridge. This
distance is related to by = n d, where n is an integer. In the schematic of Fig., d and
= 3d, and so n = 3.

LEVER simple machine for making work easier

Three factors influence a lever:

1. Load to be moved
2. Effort needed to move the load
3. Fulcrum point of balance
First-class lever

The weight and muscle act on opposite sides of the fulcrum, and in the same direction
FEffort dEffort = FLoad dLoad
Mechanical advantage = dEffort / dLoad = Fload / FEffort = 1
Second-class lever

The weight and muscle act on the same side of the fulcrum and the weight is nearer to the
fulcrum, so m < w
Mechanical advantage > 1
The force (effort) arm would always greater then the load (resistance) lever arm
Third-class lever

The weight and muscle are again on the same side of the fulcrum, but the muscle is nearer to
the fulcrum than the weight m>>w
Mechanical advantage < 1
The load (resistance) arm would always greater then the force (effort) lever arm
Example: The distance from the center of the wheel to the handles on a
wheelbarrow is 60 inches. The weight in the bucket is 18 inches from the center
of the wheel. If 300 lb is placed in the bucket, how much force must be applied
at the handles to lift the wheelbarrow?
FEffort dEffort = FLoad dLoad
FEffort 60 in = 300 Ib 18 in
FEffort = 90 Ib
The mechanical advantage of the lever in this example would be:
Mechanical advantage = 60 / 18 = 300 / 90 = 3.33
Muscle 104 106 fibers
Fiber 10 80 m diameter
Let us say 50 m typical, So there are 1 / (50 m)2= 4104 fibers/cm2
Fiber 2000 myofibrils
Myofibril 1500 myosin filaments + 3000 actin filaments ~ 1104 - 1.7105
Myosin filament 200 myosin molecules
Thank you