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    Skeletal Muscle BasicsContraction and Basic mechanical

    properties

    Taken from:Professor Bruce Lynn

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    Skeletal Muscle Basics

    3 Lectures:

    Basic structure of muscle

    Muscle activation & relaxation

    Basic mechanical properties

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    Outline

    Sliding filaments & the crossbridge cycle

    Force & Power

    Antagonistic muscles

    Series & Parallel structures

    Arrangement of fibres with muscle

    Force-velocity relation, also power

    For shortening & stretch

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    Dependence of isometric force on sarcomere length

    Sarcomere length (% of optimum)

    Force (%

    of max)

    Force is proportional to filament overlap:

    important evidence for sliding filaments

    The Tension Length Curve

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    What causes the filament sliding?

    Myosin heads bind to actin, then gothrough a cycle of events the cross

    bridge cycle

    Overall effect is force generation and ATP

    hydrolysis

    As all myosin molecules are identical, can

    reduce problem to considering just a

    single myosin head interacting with actin

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    Z

    ADP Pi

    1

    Z

    ADP Pi

    attach

    2

    Z

    ADP

    Pi release &

    weak to strong

    Pi

    3

    Z

    ADP release

    & filamentsliding

    ADP4ATP

    binding

    Z

    ATP

    ATP

    5

    detach &

    ATP

    hydrolysis

    Does not occur

    when [Ca] low

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    Force in Isometric contraction: no sliding

    Thin filament

    Thick filament

    Attached crossbridge, no

    force (spring not stretched)

    Attached crossbridge has

    changed shape to stretch

    spring, force but no sliding

    2

    3Direction of isometric force: toward M line

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    Cross-bridge Cycle: Key Features

    1)ATP is used in each cycle to provide the energyRigor mortis occurs if ATP concentration = 0

    2) Direction of filament force and sliding (if sliding

    occurs) is one-way (thin filament moves toward M-line

    at the centre of the sarcomere)

    3) Step size is small: sliding produced by one cycle is

    only about 1% of the sarcomere lengthMany cycles occur in succession to cause largemovements (as in running, walking, etc)

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    Why so complicated?

    Some constraints due

    to muscle properties

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    Outline

    Sliding filaments & the crossbridge cycle

    Force & Power

    Antagonistic muscles

    Series & Parallel structures

    Arrangement of fibres with muscle

    Force-velocity relation, also power

    For shortening & stretch

    What are they? How are they

    different?

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    Isometric

    Force

    Power

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    What is isometric contraction?

    Muscles are active (=contracting)

    producing isometric force

    The muscle force resists gravity and

    prevents the arm and book falling

    Isometric means the muscle length is

    constant

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    Contraction with shortening (concentric)

    Biceps contracts and its shortening

    flexes the elbow

    Biceps does work lifting the book

    POWER is the rate at which work

    is done

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    Outline

    Sliding filaments & the crossbridge cycle

    Force & Power

    Antagonistic muscles

    Series & Parallel structures

    Arrangement of fibres with muscle

    Force-velocity relation, also power

    For shortening & stretch

    Required due to crossbridge cycle

    & sliding filament arrangement

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    Tendon

    Tendon

    Biceps

    Tendon

    Triceps

    Tendon

    Example: Rotation around the elbow

    Active (contracting) muscle can shorten (pull towards its

    center)

    Therefore, antagonistic muscles are required

    BUT it cannot elongate (push away from its center)

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    Rotation around the elbow: Flexion

    Biceps contracts & shortens

    Triceps is lengthened

    (not contracting)

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    Rotation around the elbow: Extension

    Biceps is lengthened

    (not contracting)

    Triceps contracts & shortens

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    Outline

    Sliding filaments & the crossbridge cycle

    Force & Power

    Antagonistic muscles

    Series & Parallel structures

    Arrangement of fibres with muscle

    Force-velocity relation, also power

    For shortening & stretch

    How the arrangement ofstructures affect force and

    length change

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    Springs connected in Parallel

    Fixed position

    Springs connected in Series

    Fixed position

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    Structures in Series:

    Force at A and B are equal.

    For structures in series, forces do NOT add up

    Fixed position

    A

    Fixed position

    A B

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    A

    Fixed position

    Force = A

    Fixed position

    A

    B

    Force = A+B

    For structures in parallel, forces add up

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    Length changes

    Length 2.0 m

    Length 1.5 m

    = 0.5 m

    Length 3.0 m

    = 1.0 m

    Length 4.0 m

    Connect in series

    For structures in series, length changes add up

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    Length changes

    Length 2.0 m

    Length 1.5 m

    = 0.5 m

    Length 2.0 m

    Connect in parallel

    = 0.5 mLength 1.5 m

    For structures in parallel, length changes do NOT add up

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    Contractile and Elastic Structures

    In series and in parallel

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    A muscle-tendon complex (MTC)

    Bone

    Bone

    Muscle fibresor Contractile Component (CC)

    Tendon

    or Series elastic component (SEC)

    Parallel elastic

    component

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    A muscle-tendon complex (MTC)

    Because muscle and tendon are in series:

    Both experience the same force at each

    moment. An observed length change of MTC could

    be due to either component

    Tendon can only be stretched when

    muscle is active

    Muscle cannot move bones without first

    stretching tendon

    Bone

    Bone

    Muscle

    or CC

    Tendon

    or SEC

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    Elasticity also in parallel

    The parallel element:

    Can exert force when CC

    is relaxed.

    Adds its force to that of

    muscle when CC is active.

    More complicatedconnections can switch

    elasticity between series

    and parallel.

    Bone

    Bone

    CC

    SEC

    PEC

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    Where and what are the SEC and PEC

    relative to the crossbridges?

    Tendon (collagen) series

    Aponeuroses (collagen) series

    Epimysium (collagen) parallel

    Filaments (titin) parallel

    Filaments (myosin, actin) series

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    Outline

    Sliding filaments & the crossbridge cycle

    Force & Power

    Antagonistic muscles

    Series & Parallel structures

    Arrangement of fibres with muscle

    Force-velocity relation, also power

    For shortening & stretch

    How the arrangement ofstructures affect force and

    length change

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    Arrangement of muscle fibres: some examples

    parallel

    fusiform

    triangular

    unipennate

    bipennate

    multipennate

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    Volumes equal and line of muscle force is the same

    Each fibre in pennate muscle is half the length

    of the fibres in the parallel muscle

    and at angle to the line of muscle force;

    force along line of muscle (F) = cos * force

    along line of fibre (f)

    For = 30o, cos = 0.87

    But there are twice as many fibres in thepennate muscle as in the parallel muscle

    Net effect: pennate muscle produces 2 * 0.87 =

    1.74 times more force than the parallel muscle

    Arrangement of fibres within muscle:

    Pennation increases muscle force

    parallel pennate

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    Net effect: pennate muscle shortening is only 0.5 * 0.87

    = 0.41 times as much as the parallel muscle per unit

    time

    the cos rule means that muscle shortening

    is cos * fibre shortening.

    Pennation reduces muscle shortening velocity

    also each fibre in the pennate muscle only

    shortens half as far as each fibre in the

    parallel muscle.

    In each unit of time

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    Outline

    Sliding filaments & the crossbridge cycle

    Force & Power

    Antagonistic muscles

    Series & Parallel structures

    Arrangement of fibres with muscle

    Force-velocity relation, also power

    For shortening & stretch

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    Contraction with shortening (concentric)

    Biceps contracts and its shortening

    flexes the elbow

    Biceps does work lifting the book

    POWER is the rate at which work

    is done

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    Muscle Force

    Muscle length

    time(Lever movement)

    Before stimulation of the muscle

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    start stimulation of the muscle

    Isometric phasemuscle force toosmall to lift weight

    Muscle Force

    Muscle length

    time

    (Lever movement)

    Stim

    I i h i

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    During stimulation, muscle force

    enough to lift weight

    Muscle Force

    Muscle length

    time

    (Lever movement)

    Stim

    Isotonic shortening:constant forceduring shortening

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    Muscle Force

    Muscle length

    time(Lever movement)

    Before stimulation of the muscle

    Larger weight

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    During stimulation of the muscle

    time

    Muscle Force

    Muscle length

    (Lever movement)

    Stim

    Isometric phasemuscle force toosmall to lift weight

    I t i h t i

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    During stimulation of the muscle

    Muscle Force

    Muscle length

    time

    (Lever movement)

    Stim

    Isotonic shortening:constant forceduring shortening

    Larger force &

    slower velocity

    P k t

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    Power = work rate

    Velocity

    0.0 0.5 1.0 1.5

    Power

    0.0

    0.1

    0.2

    Velocity

    0.0 0.5 1.0 1.5

    Force

    0.0

    0.5

    1.0

    Inverse relation between

    force and velocity of

    shortening

    The Force Velocity Curve

    = (force x length ) / time

    = force x (length / time)

    = force x velocity

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    Contraction with lengthening (eccentric)

    Biceps is acting as a brake.

    Biceps is producing force, EMG, etc,(=contracting)

    The book is lowered in a slow,controlled movement.

    The elbow extends as the length of

    biceps increases due to the book &

    gravity. Work is done on biceps.

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    Force during isovelocity stretch of active

    muscle

    stim

    stim

    stim

    Force-Velocity relation for Stretch

    Velocity

    stretch shorten

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    Stretch of active muscle

    Occurs during normal every-day activities

    Contracting muscle fibres act as a brake

    Large forces can be produced

    But not much fuel (ATP) is used

    Forces can be large enough to cause

    damage

    Not covered in many standard textbooks

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    Skeletal Muscle BasicsContraction. Basic mechanical properties

    Summary

    Tension-length curve, max force at max filament overlap

    Cross bridge cycle, myosin head binds to actin, ATP splitting,

    repetitiveMuscle morphology:

    short fat muscles, high force, low speed;

    long thin muscles, low force, high speed

    Inverse force-velocity relationPower = Force*velocity; max power at ca 1/3 max force or velocity

    Eccentric contractions, high force.

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    Skeletal Muscle Basics

    Contraction. Basic mechanical properties

    Good source of information

    Jones et al., Skeletal Muscle from Molecules to Movement, 2004,

    Churchill Livingstone.

    Acknowledgements

    Thanks for Nancy Curtin, Imperial College, for use of many of her slides.