Introduction to Biomechanics Assigned Reading: Jenkins: pp. 17-30. Definitions biomechanics - application of the principles of mechanics / physics to biological organisms muscle action vs. function o 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 o function - how the organism chooses to use a muscle - can involve positive, negative or non- work (see below) 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 o - occurs when a force moves its point of application o - e.g., muscles work by moving myosin heads along actin filament o - 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 o - work is measured in joules [J = (N)(m)] o - 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 o - limited to experimental conditions in which mechanical properties (either tension or change in length) of muscle are measured; i.e., not possible in vivo o - isometric contraction - muscle length is fixed and tension is measured - used to generate length/tension curves (see below) o - 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 o - theory proposed by biophysicist Jean Hanson (1919-73) and physiologist Hugh Esmor Huxley (1924- ) in 1954 o - states that during contraction thin filaments slide past thick filaments with no change in the length of either type of filament o - force for producing sliding of thin filaments is generated by the crossbridges (formed by myosin heads) o - 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: o 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 o 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 o - generated from series of isotonic contractions o - 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 o - 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 o - specifically F = total CSA x Specific Tension of muscle (N/cm2) o - 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 o - maximum sarcomere excursion = 50% of resting length o - thus longer fibers will contract a greater distance 3. Velocity (distance of shortening/unit of time) is proportional to fiber length (assuming equal load) o - muscles of different fiber length will contract to 50% in same amount of time o - 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 o - variation in muscle architecture can affect: (1) excursion (distance a muscle can contract) (2) velocity (3) force, and (4) line of action o - variety of classification schemes exist; none perfect (except mine); many primarily descriptive o - functionally 3 general types: parallel, triangular and pinnate based on fiber arrangement 1) triangular - muscle fibers radially arranged -specialized for altering line of action assuming nonuniform 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 Advantage of pinnation / Disadvantage of parallel o - maximum force produced by a muscle is proportional to the sum of the cross-section of all its fibers o - for muscles of equal volume, more muscle fibers can be packed into a pinnate arrangement than a parallel arrangement o - 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 o - thus force output = # of fibers x cosine of angle of insertion o - 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 o - excursion = length a muscle fiber can contract; function of fiber length o - for muscles of equal length, pinnate muscles have decreased excursion relative to parallel o o o o - 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 o - machine - device for transmitting forces from one point to another o - majority (but not all) of skeletal elements function as type of machine known as lever o - lever is a rigid bar (regardless of shape) which rotates about a fixed point (fulcrum) o - 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 Lever systems are most easily analyzed under the conditions of equilibrium Force equilibrium: Fi x Li (in-torque) = Fo x Lo (out-torque) o - 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 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 Note that for a given muscle input (Fi) a muscle lever system can either: o o 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 Muscle fiber types Quality of force production can be varied by using different types of muscle fibers o - vertebrate muscles can be broadly divided into slow and fast based upon speed of contraction o - 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 o - fast [twitch] 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 1) mosaic - fast and slow fibers uniformly distributed 2) compartmentalized - fiber types non-uniformly distributed into intramuscular compartments o - however, some muscles which are used for repetitive or constant tasks (e.g., posture) can be comprised nearly entirely of slow fibers - e.g., soleus