Musculoskeletal 4: Mechanical properties of skeletal muscle

Norsila Abdul Wahab Department of Physiology 29th August 2006

Terms
Muscular Strength - ability of a given muscle group to generate torque at a particular joint Muscular Power - product of force and velocity Muscular Endurance - ability of muscle group to exert tension over time

Characteristics of muscles
Muscles are contractile tissues with four characteristics
Electrical excitability responds to stimuli Contractility able to shorten in length Extensibility stretches when pulled Elasticity return to original shape and length after contraction

Muscle twitch
A brief contraction followed by relaxation (caused by a single AP) The twitch starts about 2ms after the start of depolarization, before repolarization is complete Duration of twitch varies with muscle type

Fiber Types
Type I
Slow twitch Red muscles For long, slow, posture-maintaining contractions Eg long muscles of the back

Type II
Fast twitch White muscles For fine, skilled movement Eg extraocular muscles, some hand muscles

The response of a muscle to a single stimulus = muscle twitch The force developed = twitch tension Intensity of stimulus increased > more motor fiber will be recruited > increased in number of active muscle fibers and the total tension developed

MECHANICAL PROPERTIES OF SKELETAL MUSCLE

Amount of force a muscle exerts depends on
No. of active muscle fibers Frequency of stimulation Rate at which the muscle shortens Initial resting length of muscle Cross-sectional area of the muscle The length-tension relationship The force-velocity relationship Summation of contractions in single muscle fibers Nature (type) of contraction
isotonic, Isometric eccentric contractions

Motor unit summation and graded muscle responses

The length-tension relationship

In general, the maximum amount of tension is generated when the muscle is near resting length When the muscle is near resting length, the maximum possible number of myosin crossbridges can be formed. When a muscle is shortened too much, the ends of the thin actin filaments from opposite ends of the sarcomere begin to interfere with one another.

Length-tension relationship

The length-tension relationship

If the muscle contracts too much, actin from one end of the sarcomere may bind with myosin at the far end of the sarcomere, which would pull the actin in the wrong direction. When a muscle is overstretched, the actin filaments are pulled away from many of the myosin heads - far fewer actin-myosin cross-bridges can form and less power can be generated.

Degree of actin and myosin filament overlap effect on tension

At point C
actin has pulled all the way out nearer to the end of the myosin filament The tension developed by the activated muscle is near zero

Degree of actin and myosin filament overlap effect on tension

B

B*

As sarcomere shortens and actin begins to overlap the myosin filament, the tension increases progressively until the sarcomere length decreases to about 2.2 micrometers.

Degree of actin and myosin filament overlap effect on tension

Point B
Actin filament has overlapped all the cross-bridges of the myosin filament Upon further shortening the sarcomere maintains full tension until point B* at approx 2.0 micrometers

Length-tension relationship
Maximum contraction occurs when there is maximum overlap between actin and the cross-bridges of the myosin The greater the number of cross-bridges pulling the actin filaments, the greater the strength of contraction

The ends of two actin filaments begin to overlap each other, in addition to overlapping the myosin filaments Point A
the strength of contraction decreases The two Z discs abut the ends of the myosin filaments

As contraction proceeds to shorter sarcomere lengths, the ends of the myosin filaments are crumpled, strength of contraction decreases

The force-velocity relationship

The greater the load on the muscle, the slower the muscle will be able to contract. The multiple force velocity curves depicted also demonstrate that, when a muscle is moved away from resting length, it contracts with slightly less speed and it cannot handle as great a load. ie, a muscle contracts fastest, and most powerfully, when it is near resting length.

Force-velocity curve

Summation of contractions in single muscle fibers

If a muscle is activated by a pair of stimuli that are so close together that the second stimulus arises before the muscle has fully relaxed after the first, the total tension developed following the second stimulus is greater than that developed in response to a single stimulus

Summation of contractions in single muscle fibers

Repeated activation of motor neurons at a sufficiently high frequency results in summation of the contractile force of individual muscle twitches. This occurs when stimulation (AP) frequency is too high to allow calcium ions to be resequestered by the SR between muscle twitches. Summation of high frequency stimuli results in smooth, coordinated muscle contractions. This phenomenon is known as "summation of twitches"

Tetanus
If a number of stimuli are given in quick succession, the tension developed progressively summates and this response = tetanus

Twitch and tetanus

Motor unit summation and graded muscle responses

Muscle fibers within individual motor units contract in an all-or-none manner. How do muscle get the additional contractile force (graded muscle response)?
Recruiting additional motor units during a muscle contraction Motor units are recruited in order of size, beginning with the smallest and leading up to the largest motor unit. Motor cortex activates motor units in sequence, allowing some motor units to relax while others are performing the work. This "cycling" between activation of motor units gives the motor units being used brief rest periods so that they can work for extended periods of time.

Treppe (staircase phenomenon)

If the voltage of electrical shocks delivered to an isolated muscle in vitro is gradually increased from zero, the strength of the muscle twitches will increase accordingly, up to a maximal value where all fibers are stimulated. Graded nature of muscle contraction If these series of shocks is given to a fresh muscle so that each shock produces a separate twitch, each of the twitches will be successively stronger, up to a higher maximum -> treppe effect Treppe may represent a warm up effect, believed to be due to an increased in intracellular calcium ions

Types of contraction
Isometric Isotonic Eccentric

Isometric contractions
A contraction in which the muscle length remains constant as the tension in the muscle increases. Cross-bridge formation between actin and myosin is capable of increasing the tension within the muscle (muscle tone) while the work being done prevents the muscle length from decreasing. An example of isometric contraction would be the increase in tension you feel in your muscles when you push against an immovable object like a wall.

Isotonic contractions
Muscle is shortening but the tension remains constant. Cross-bridge formation and cycling between cross-bridges is leading to the shortening of the muscle The muscle is shortening and may or may not be moving an object.

Eccentric contractions
Eccentric contraction is very similar to isotonic contraction except that instead of shortening, the muscle is lengthening during the contraction. Most normal physical activities involve a combination of isometric, isotonic and eccentric contractions.