Definitions
- Bio mechanics – application of the principles of mechanics / physics to biological organisms
- muscle action vs. function
- action – motions produced by a muscle’s shortening; described in reference to axes and planes of body
- – determined by architecture of joint and position of muscle around it
- – pure mechanics which can be inferred
- function – how the organism chooses to use a muscle
- – can involve positive, negative or non- work (see below)
- action – motions produced by a muscle’s shortening; described in reference to axes and planes of body
- agonists (Gr., contestant)– muscles with an identical action; usually restricted to single axis or plane of reference
- antagonists (Gr., against + contestant; lit. enemy) – muscles with opposite action; usually restricted to single axis or plane of reference
- synergist (Gr., together + work) – muscles which act together to perform a function; can involve both agonists and antagonists
- work (W) – the mechanical definition
- – occurs when a force moves its point of application
- – e.g., muscles work by moving myosin heads along actin filament
- – thus W = (F) (X), where
- F= force (newtons; N)
- X = distance moved (meters; m); can be positive or negative relative to line of action of force
- – work is measured in joules [J = (N)(m)]
- – muscles generate tension to perform positive, negative or non- work
- – positive work – muscle shortens while generating tension (i.e., X > 0)
- – negative work – muscle lengthens while generating tension (i.e., X < O)
- – non-work – muscle generates tension without changing length (X=0)
Some Physiology (gag!)
- isometric and isotonic contractions
- – limited to experimental conditions in which mechanical properties (either tension or change in length) of muscle are measured; i.e., not possible in vivo
- – isometric contraction – muscle length is fixed and tension is measured
- – used to generate length/tension curves (see below)
- – isotonic contraction – muscle tension (load) is fixed and change in length (shortening) measured
- – used to generate force (tension)/velocity curves (see below)
- sliding filament theory and length/tension curve (see Figure 1)
- – theory proposed by biophysicist Jean Hanson (1919-73) and physiologist Hugh Esmor Huxley (1924- ) in 1954
- – states that during contraction thin filaments slide past thick filaments with no change in the length of either type of filament
- – force for producing sliding of thin filaments is generated by the cross-bridges (formed by myosin heads)
- – theory predicts that force output will be proportional to the degree of overlap between thick and thin filaments or, more specifically, the number of cross-bridges formed
- biomechanical implications of the sliding filament theory:
- 1. for maximum force output (total tension) muscle should be positioned below its optimal length so that work (either positive or negative) will occur over peak of length/tension curve
- 2. muscles which produce the same action across a joint are typically arranged such that their optimal lengths occur at different joint positions thus permitting a nearly constant level of force output at all joint positions
- velocity-force curves (see Figure 2)
- – generated from series of isotonic contractions
- – force and velocity are inversely related such that at zero (0) velocity maximum force is generated, and at maximum velocity zero (0) force is generated
- – power output = force x velocity (rate of doing work)
- – measured in watts (1N x 1m/s)
- – is maximized at about 30% of maximum force
Preliminary concepts
- 1. Force output is proportional to cross-sectional area (see Figure 3)
- – specifically F = total CSA x Specific Tension of muscle (N/cm2)
- – thus muscles that differ in length but have equal CSA generate equal amounts of force
- 2. Excursion (distance a muscle can shorten) is proportional to fiber length (see Figure 4)
- – maximum sarcomere excursion = 50% of resting length
- – thus longer fibers will contract a greater distance
- 3. Velocity (distance of shortening/unit of time) is proportional to fiber length (assuming equal load)
- – muscles of different fiber length will contract to 50% in same amount of time
- – since excursions distances differ but time is constant, velocity is greater in muscles with longer fibers
Muscle Architecture
- Muscle architecture refers to arrangement and length of muscle fibers w/i a muscle
- – variation in muscle architecture can affect:
- (1) excursion (distance a muscle can contract)
- (2) velocity
- (3) force, and
- (4) line of action
- – variety of classification schemes exist; none perfect (except mine); many primarily descriptive
- – functionally 3 general types: parallel, triangular and pinnate based on fiber arrangement (see Figure 5)
- 1) triangular – muscle fibers radially arranged
- -specialized for altering line of action assuming non-uniform distribution of motor units
- 2) parallel – muscle fibers are arranged parallel to line of action (muscle pull)
- – specialized for excursion and/or velocity
- 3) pinnate – muscle fibers lie at an angle to line of action (muscle pull)
- – specialized for force production
- – N.B. Relationship between angle of pinnation (parallel fibers have an angle of pinnation = 0 degrees), fiber length and excursion is not simple; in fact in some situations pinnation actually can increase excursion
- 1) triangular – muscle fibers radially arranged
- – variation in muscle architecture can affect:
- Advantage of pinnation / Disadvantage of parallel (see Figure 6)
- – maximum force produced by a muscle is proportional to the sum of the cross-section of all its fibers
- – for muscles of equal volume, more muscle fibers can be packed into a pinnate arrangement than a parallel arrangement
- – since axis of contraction of muscle fibers not parallel to pull of muscle (line of action) some muscle force dissipated perpendicular to line of action
- – thus force output = # of fibers x cosine of angle of insertion
- – thus advantage of pinnation is to increase force output of a muscle by packing more fibers in a given volume of space
- Cost of pinnation / Advantage of parallel (see Figure 7)
- – excursion = length a muscle fiber can contract; function of fiber length
- – for muscles of equal length, pinnate muscles have decreased excursion relative to parallel
- – max. sarcomere shortening = 50% of resting length; thus max. excursion of muscle = 50% of fiber length
- – parallel fibers can shorten to their maximum
- – pinnate fibers cannot shorten to their maximum w/o dislodging themselves from their tendons
- – thus pinnate muscle has shorter excursion
Lever mechanics
- Muscles generate forces and skeletal elements apply these forces and thus serve a machines
- – machine – device for transmitting forces from one point to another
- – majority (but not all) of skeletal elements function as type of machine known as lever
- – lever is a rigid bar (regardless of shape) which rotates about a fixed point (fulcrum)
- – in levers forces work by creating rotational forces about the joints (fulcrum) known as moments; i.e.,
- m = F x L, where
- m = moment or torque
- F = force; in this case muscle tension
- L = Lever (or moment) arm; distance between force and fulcrum; lies perpendicular to line of action of force
- m = F x L, where
- Lever systems are most easily analyzed under the conditions of equilibrium (see Figure 8)
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- Force equilibrium: Fi x Li (in-torque) = Fo x Lo (out-torque)
- – solving for Fo:
- Fo = Fi x (Li/Lo)
- – thus to maximize force-output of a lever system for a given muscle force (Fi):
- 1) increase Li
- 2) decrease Lo
- – Li/Lo = lever advantage
- – solving for Fo:
- Force equilibrium: Fi x Li (in-torque) = Fo x Lo (out-torque)
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- Velocity equilibrium: Vo x Li = Vi x Lo
- – solving for Vo
- Vo = Vi x (Lo/Li)
- – thus to maximize velocity-output of a lever system for a given muscle velocity (Vi):
- 1) decrease Li
- 2) increase Lo
- – Lo/Li = gear ratio
- – solving for Vo
- Velocity equilibrium: Vo x Li = Vi x Lo
- Note that for a given muscle input (Fi) a muscle lever system can either:
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- a) maximize lever advantage (Li/Lo) and produce a stronger but slower force (Fo)
- b) maximize gear ratio (Lo/Li) and produce a faster but weaker force (Vo)
- – it cannot maximize both (inverse relationship)
- – thus, there is a trade off between velocity and force in any lever system
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Muscle fiber types
- Quality of force production can be varied by using different types of muscle fibers
- – vertebrate muscles can be broadly divided into slow and fast based upon speed of contraction
- – slow fibers – specialized for prolonged tension generation
- – typically generate small forces (due to small fiber CSA and low innervation ratio) at low metabolic cost (aerobic respiration)
- – fatigue resistant due to high density of mitochondria and myoglobin
- – 2 subtypes
- 1) tonic – multi-terminal fibers; membrane cannot propagate an AP thus contraction is graded; limited to extra-ocular muscles in mammals
- 2) slow twitch – single terminal fibers; widely distributed
- – fast fibers – specialized for generating tension rapidly
- – typically generate larger forces (due to larger fiber CSA and high innervation ratio) at high metabolic cost (use both aerobic and anaerobic respiration)
- – different sub-types (2A, 2B, 2X) differ in myosin isoforms and fatigue resistance
- Majority of muscles are of mixed fiber type composition being a combination of fast and slow fibers occurring in two arrangements
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- 1) mosaic – fast and slow fibers uniformly distributed
- 2) compartmentalized – fiber types non-uniformly distributed into intramuscular compartments
- – however, some muscles which are used for repetitive or constant tasks (e.g., posture) can be comprised nearly entirely of slow fibers
- – e.g., soleu
THANKS TO FLORIDA INTERNATIONAL UNIVERSITY
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