KIN 840

2006-1

Muscle Mechanics

2006 by Stephen Robinovitch, All Rights Reserved

Outline
anatomical structure eccentric and concentric contractions determinants of muscle force (stimulation frequency, muscle bre type, muscle length and velocity, crosssectional area) size principle Hill's equation for muscle shortening Hills active state model of muscle contraction quick-release experiments Huxley's sliding lament model Loeb's "virtual muscle" model

Muscle anatomy

Eccentric and concentric contractions

Concentric (energy generating, positive work) contractions tend to increase joint angular velocity, and increase the total energy of the system Eccentric (energy absorbing, negative work) contractions tend to decrease (or brake) joint angular velocity, and reduce the total energy of the system
Concentric (accelerating) contraction of m2 Eccentric (braking) contraction of m2

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
Joint moment (Nm) extensor

B3 A3 B1 A1 B2 A2
exion

A4 B4

extension

B7 A7
exor

A6 B6 A5 B5

Joint rotation (radians)

Muscle force results from interaction between contractile proteins

sarcomere: the smallest anatomical unit that contracts like a muscle sliding lament model proposes that muscle force arises from cyclic binding between thick and thin laments of the sarcomere thin laments contain actin, troponin C, and tropomyosin thick laments 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 myobrils) 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 muscle force development (a partial list)

muscle bre type number of activated motor neurons, frequency of discharge muscle length velocity of shortening/ lengthening muscle geometry (physiological cross-sectional area (PCSA), angle of pennation)

Muscle-nerve interaction
a motor nerve enters muscle and splits into numerous axons; each axon contacts 102000 muscle bres each muscle bre 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 bres it contacts

muscle platysma Brachioradialis Tibialis anterior gastrocnemius

# muscle fibers 27,100 130,000 250,000 1,120,000

# motor units 1,100 330 450 580

av. fibers per motor u. 25 410 600 2,000

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 bres and motor units, dened by contraction speed, peak force, fatigue resistance

Size Principle
When a stimulus is applied to the ventral aspect of the spinal cord, the smallest and most excitable motor units are activated rst. These tend to be slow (S) motor units which innervate slow oxidative (SO) muscle bres. Larger FR and FF motor units that innervate FOG and FG bres are recruited only at high levels of force. Sequence is reversed when force level falls, with largest motor units dropping out rst.

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 laments reduces ability of crossbridges to develop tension left of (D): myosin laments collide with Z-lines and fold, and force declines rapidly

Muscle length affects force development 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

Muscle force-velocity behaviour is described by the Hill Equation

An empirical relation that describes the force-velocity behaviour of muscle during shortening is the Hill Equation.

Greatest force is developed when lengthening near resting length

Hills active state model of muscle contraction

Hill assumed: (1) for a given length, muscle always develops the same peak force T0(x1,t); (2) if the muscle is shortening, some force is dissipated in overcoming inherent viscous resistance B: muscle damping constant, which must be a nonlinear function of shortening velocity and temperature KSE: stiffness of the series elastic component; represents forcedeection properties of tendon KPE: stiffness of the parallel elastic component; represents forcedeection properties of sarcolemma, epimysium, perimysium, and endomysium

Quick-release experiments for determining the Hill model parameters

hold muscle length xed with the catch stimulate muscle to produce peak (isometric) force T0 instantly release catch 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)

there is an instant change (! x2) in the length of KSE following release this is followed by a more gradual change (!x1) in the length of the muscle as T increases, there is a decrease in v (slope of dashed line), reecting that muscle cannot shorten quickly under high loads combinations of T and v reect the force-velocity properties of a given muscle

Shortcomings of the active state model

the 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 the model 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)

Huxleys sliding lament 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

Gerry Loebs virtual muscle (J Neuroscience

Methods, 2000)
each muscle consists of (1) a contractile element, (2) a series elastic element, and (3) a muscle mass instabilities in the model are prevented by the muscle mass, and a small viscosity in the parallel elastic element contractile element consists of several motor units (3-5 for each ber type), each dened by its order of recruitment, ring frequency, force-lengthvelocity relationships, passive parallel elastic element, and total forceproducing capacity (proportional to total crosssectional area)
Matlab/ Simulink-based model for muscle and tendon distributed freely over the Internet at http://ami.usc.edu/projects/ami/projects/bion/muscu loskeletal/virtual_muscle.html

Virtual Muscle (cont)

the motor nucleus of the whole muscle receives a single, time-varying neural activation command signal, which activates each motor unit according to the userdened recruitment order within each motor unit, the frequency of motoneuronal ring is modulated in a realistic manner the activation signal can be based on pre-recorded EMG data, simulated feedback-driven reex dynamics, or cortical commands

Virtual muscle (cont)

user denes the properties of each ber type and the morphometry of each musculotendon element, including the elastic properties of tendon and aponeurosis user then integrates these elements with their dened skeletal dynamics and control system (developed in Working Model)