muscle
TRANSCRIPT
Required readings: Biomechanics and Motor Control of Human
Movement (class text) by D.A. Winter, pp. 165-212
Next Class• Reading assignment
– Biomechanics of Skeletal Muscle by T. Lorenz and M. Campello (adapted from M. I. Pitman and L. Peterson; pp. 149-171
– EMG by W. Herzog, A. C. S. Guimaraes, and Y. T. Zhang; pp. 308-336
– http://www.delsys.com/library/tutorials.htm• Surface Electromyography: Detecting and Recording• The Use of Surface Electromyography in Biomechanics
• Exam on anthropometry• Turn in EMG abstract• Prepare short presentation on EMG research article• Laboratory experiment on EMG• Hour assigned
Advanced Biomechanics of Physical Activity (KIN 831)
Muscle – Structure, Function, and Electromechanical Characteristics
•Material included in this presentation is derived primarily from two sources: Jensen, C. R., Schultz, G. W., Bangerter, B. L. (1983). Applied kinesiology and biomechanics. New York: McGraw-Hill Nigg, B. M. & Herzog, W. (1994). Biomechanics of the musculo-skeletal system. New York: Wiley & Sons Nordin, M. & Frankel, V. H. (1989). Basic Biomechanics of the Musculoskeletal System. (2nd ed.). Philadelphia: Lea & Febiger Winter, D.A. (1990). Biomechanical and motor control of human movement. (2nd ed.). New York: Wiley & Sons
Introduction• Muscular system consists of three muscle
types: cardiac, smooth, and skeletal
• Skeletal muscle most abundant tissue in the human body (40-45% of total body weight)
• Human body has more than 430 pairs of skeletal muscle; most vigorous movement produced by 80 pairs
Introduction (continued)
• Skeletal muscles provide strength and protection for the skeleton, enable bones to move, provide the maintenance of body posture against gravity
• Skeletal muscles perform both dynamic and static work
Muscle Structure• Structural unit of skeletal muscle is the
multinucleated muscle cell or fiber (thickness: 10-100 m, length: 1-30 cm
• Muscle fibers consist of myofibrils (sarcomeres in series: basic contractile unit of muscle)
• Myofibrils consist of myofilaments (actin and myosin)
Microscopic-Macroscopic Structure of
Skeletal Muscle
Muscle Structure (continued)• Composition of sarcomere
– Z line to Z line ( 1.27-3.6 m in length)– Thin filaments (actin: 5 nm in diameter)– Thick filaments (myosin: 15 nm in diameter)– Myofilaments in parallel with sarcomere– Sarcomeres in series within myofibrils
Muscle Structure (continued)• Motor unit
– Functional unit of muscle contraction– Composed of motor neuron and all muscle cells
(fibers) innervated by motor neuron– Follows “all-or-none” principle – impulse from
motor neuron will cause contraction in all muscle fibers it innervates or none
•Smallest MU recruited at lowest stimulation frequency
•As frequency of stimulation of smallest MU increases, force of its contraction increases
•As frequency of stimulation continues to increase, but not before maximum contraction of smallest MU, another MU will be recruited
•Etc.
Size Principle• Smallest motor units recruited first
• Smallest motor units recruited with lower stimulation frequencies
• Smallest motor units with relatively low levels of tension provide for finer control of movement
• Larger motor units recruited later with increased frequency of stimulation and increased need for greater tension
Size Principle
• Tension is reduced by the reverse process– Successive reduction of firing rates– Dropping out of larger units first
Muscle Structure (continued)
• Motor unit– Vary in ratio of muscle fibers/motor neuron
• Fine control – few fibers (e.g., muscles of eye and fingers, as few as 3-6/motor neuron), tetanize at higher frequencies
• Gross control – many fibers (e.g., gastrocnemius, 2000/motor neuron), tetanize at lower frequencies
– Fibers of motor unit dispersed throughout muscle
•Motor Unit•Tonic units – smaller, slow twitch, rich in mitochondria, highly capillarized, high aerobic metabolism, low peak tension, long time to peak (60-120ms)
•Phasic units – larger, fast twitch, poorly capillarized, rely on anaerobic metabolism, high peak tension, short time to peak (10-50ms)
Muscle Structure (continued)
• Motor unit (continued)– Weakest voluntary contraction is a twitch
(single contraction of a motor unit)– Twitch times for tension to reach maximum
varies by muscle and person– Twitch times for maximum tension are shorter
in the upper extremity muscles (≈40-50ms) than in the lower extremity muscles (≈70-80ms)
Motor Unit Twitch
Shape of Graded Contraction
Shape of Graded Contraction• Shape and time period of voluntary tension curve
in building up maximum tension– Due to delay between each MU action potential and
maximum twitch tension– Related to the size principle of recruitment of motor
units– Turn-on times ≈ 200ms
• Shape and time period of voluntary relaxation curve in reducing tension– Related to shape of individual muscle twitches– Related to the size principle in reverse– Due to stored elastic energy of muscle– Turn-off times ≈ 300ms
Force Production – Length-Tension Relationship
• Force of contraction in a single fiber determined by overlap of actin and myosin (i.e., structural alterations in sarcomere) (see figure)
• Force of contraction for whole muscle must account for active (contractile) and passive (series and parallel elastic elements) components
Parallel Connective Tissue• Parallel elastic component
• Tissues surrounding contractile elements
• Acts like elastic band
• Slack when muscle at resting length of less
• Non-linear force length curve
• Sarcolemma, endomysium, perimysium, and epimysium forms parallel elastic element of skeletal muscle
Series Elastic Tissue• Tissues in series with contractile component• Tendon forms series elastic element of
skeletal muscle• Endomysium, perimysium, and epimysium
continuous with connective tissue of tendon• Lengthen slightly under isometric
contraction (≈ 3-7% of muscle length)• Potential mechanism for stored elastic
energy (i.e., function in prestretch of muscle prior to explosive concentric contraction)
Isometric Contraction
Musculotendinous Unit
• Tendon and connective tissues in muscle (sarcolemma, endomysium, perimysium, and epimysium) are viscoelastic
• Viscoelastic structures help determine mechanical characteristics of muscles during contraction and passive extension
Musculotendinous Unit (continued)
• Functions of elastic elements of muscle– Keep “ready” state for muscle contraction– Contribute to smooth contraction– Reduce force buildup on muscle and may
prevent or reduce muscle injury– Viscoelastic property may help muscle absorb,
store, and return energy
Muscle Model
Force Production – Gradation of Contraction
• Synchronization (number of motor units active at one time) – more force potential
• Size of motor units – motor units with larger number of fibers have greater force potential
• Type of motor units – type IIA and IIB force potential, type I force potential
Force Production – Gradation of Contraction (continued)
• Summation – increase frequency of stimulation, to some limit, increases the force of contraction
Force Production – Gradation of Contraction (continued)
• Size principle – tension increase– Smallest motor units recruited first and largest last
• Increased frequency of stimulation force of contraction of motor unit
• Low tension movements can be achieved in finely graded steps
• Increases frequency of stimulation recruitment of additional and larger motor units
• Movements requiring large forces are accomplished by recruiting larger and more forceful motor units
• Size principle – tension decrease– Last recruited motor units drop out first
Types of Muscle ContractionType of Contraction Definition Work
Concentric Force of muscle contraction resistance
Positive work; muscle moment and angular velocity of joint in same direction
Eccentric Force of muscle contraction resistance
Negative work; muscle moment and angular velocity of joint in opposite direction
Isokinetic Force of muscle contraction = resistance; constant angular velocity; special case is isometric contraction
Positive work; muscle moment and angular velocity of joint in same direction
Isometric Force of muscle contraction resistance; series elastic component stretch = shortening of contractile element (few to 7% of resting length of muscle)
No mechanical work; physiological work
Force Production – Length-Tension Relationship
• Difficult to study length-tension relationship– Difficult to isolate single agonist – Moment arm of muscle changes as joint angle
changes– Modeling may facilitate this type of study
Force Production – Load-Velocity Relationship
• Concentric contraction (muscle shortening) occurs when the force of contraction is greater than the resistance (positive work)
• Velocity of concentric contraction inversely related to difference between force of contraction and external load
• Zero velocity occurs (no change in muscle length) when force of contraction equals resistance (no mechanical work)
Force Production – Load-Velocity Relationship
• Eccentric contraction (muscle lengthening) occurs when the force of contraction is less than the resistance (negative work)
• Velocity of eccentric contraction is directly related to the difference between force of contraction and external load
Force Production – Force-Time Relationship
• In isometric contractions, greater force can be developed to maximum contractile force, with greater time
• Increased time permits greater force generation and transmission through the parallel elastic elements to the series elastic elements (tendon)
• Maximum contractile force may be generated in the contractile component of muscle in 10 msec; transmission to the tendon may take 300msec
3-D Relationship of Force-Velocity-Length
3-D Relationship of Force-Velocity-Length
Effect of Muscle Architecture on Contraction
• Fusiform muscle– Fibers parallel to long axis of muscle– Many sarcomeres make up long myofibrils– Advantage for length of contraction– Example: sartorius muscle– Force of contraction along long axis of muscle
of force of contraction of all muscle fibers– Tends to have smaller physiological cross
sectional area
(see figure)
Fusiform Fiber Arrangement
Fa
Fa = force of contraction of muscle fiber parallel to longitudinal axis of muscle
Fa = sum of all muscle fiber contractions parallel to long axis of muscle
Effect of Muscle Architecture on Contraction (continued)
• Pennate muscle– Fibers arranged obliquely to long axis of
muscle (pennation angle)– Uni-, bi-, and multi-pennate– Advantage for force of contraction– Example: rectus femoris (bi-pennate)– Tends to have larger physiological cross
sectional area
Pennate Fiber Arrangement
FmFa
Fa = force of contraction of muscle fiber parallel to longitudinal axis of muscle
Fm = force of contraction of muscle fiber
= pennation angle
Fa = (cos )(Fm)
Fa = sum of all muscle fiber contractions parallel to long axis of muscle
Effect of Muscle Architecture on Contraction (continued)
• Force of muscle contraction proportional to physiological cross sectional area (PCSA); sum of the cross sectional area of myofibrils
• Velocity and excursion (working range or amplitude) of muscle is proportional to length of myofiblril
Muscle Fiber TypesType I
Slow-Twitch Oxidative (SO)
Type IIA
Fast-Twitch Oxidative-
Glycolytic (FOG)
Type IIB
Fast-Twitch Glycolytic (FG)
Speed of contraction
Slow Fast Fast
Primary source of ATP production
Oxidative phosphorylation
Oxidative phosphorylation
Anaerobic glycolysis
Glycolytic enzyme activity
Low Intermediate High
Capillaries Many Many Few
Myoglobin content High High Low
Glycogen content Low Intermediate High
Fiber diameter Small Intermediate Large
Rate of fatigue Slow Intermediate Fast
Muscle Fiber Types (continued)• Smaller slow twitch motor units are characterized
as tonic units, red in appearance, smaller muscle fibers, fibers rich in mitochondria, highly capillarized, high capacity for aerobic metabolism, and produce low peak tension in a long time to peak (60-120ms).
• Larger fast twitch motor units are characterized as phasic units, white in appearance, larger muscle fibers, less mitochondria, poorly capillarized, rely on anaerobic metabolism, and produce large peak tensions in shorter periods of time (10-50ms).
Muscle Fiber Types (continued)• Nerve innervating muscle fiber determines
its type; possible to change fiber type by changing innervations of fiber
• All fibers of motor unit are of same type
• Fiber type distribution in muscle genetically determined
• Average population distribution:– 50-55% type I– 30-35% type IIA– 15% type IIB
Muscle Fiber Types (continued)
• Fiber composition of muscle relates to function (e.g., soleus – posture muscle, high percentage type I)
• Muscles mixed in fiber type composition
• Natural selection of athletes at top levels of competition
Electrical Signals of Muscle Fibers• At rest, action potential of muscle fiber -
90 mV;caused by concentrations of ions outside and inside fiber (resting state)
• With sufficient stimulation, potential inside cell raised to 30-40 mV (depolarization); associated with transverse tubular system and sarcoplasmic reticulum; causes contraction of fiber
• Return to resting state (repolarization)• Electrical signals from the motor units
(motor unit action potential, muap) can be recorded (EMG) via electrodes