Skeletal Muscle Basics

Contraction and Basic mechanical properties

Taken from:

Professor Bruce Lynn

Skeletal Muscle Basics 3 Lectures:

Basic structure of muscle Muscle activation & relaxation Basic mechanical properties

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

Force is proportional to filament overlap: important evidence for sliding filaments

Dependence of isometric force on sarcomere length

Force (% of max)

Sarcomere length (% of optimum)

The Tension Length Curve

What causes the filament sliding?

Myosin heads bind to actin, then go through 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

attach
Does not occur when [Ca] low
ADP Pi

Pi release & weak to strong

Pi

Z Z 3

ADP Pi ADP

detach & ATP hydrolysis

Z 5

ATP binding

ADP

ATP ATP

ADP release & filament sliding

Force in Isometric contraction: no sliding

2 Thin filament Attached crossbridge, no force (spring not stretched) Thick filament

Direction of isometric force: toward M line

Attached crossbridge has changed shape to stretch spring, force but no sliding

Cross-bridge Cycle: Key Features

1) ATP is used in each cycle to provide the energy Rigor 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 length

Many cycles occur in succession to cause large movements (as in running, walking, etc)

Why so complicated?

Some constraints due to muscle properties

Outline
Sliding filaments & the crossbridge cycle

Force & Power

What are they? How are they different?

Antagonistic muscles

Series & Parallel structures

Arrangement of fibres with muscle

Force-velocity relation, also power

For shortening & stretch

Isometric Force

Power

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

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

Outline
Sliding filaments & the crossbridge cycle

Force & Power

Antagonistic muscles
Required due to crossbridge cycle & sliding filament arrangement

Series & Parallel structures

Arrangement of fibres with muscle

Force-velocity relation, also power

For shortening & stretch

Active (contracting) muscle can shorten (pull towards its center) BUT it cannot elongate (push away from its center) Therefore, antagonistic muscles are required

Example: Rotation around the elbow

Tendon Biceps Tendon

Tendon

Triceps
Tendon

Rotation around the elbow: Flexion

Biceps contracts & shortens

Triceps is lengthened (not contracting)

Rotation around the elbow: Extension

Biceps is lengthened (not contracting) Triceps contracts & shortens

Outline
Sliding filaments & the crossbridge cycle

Force & Power

Antagonistic muscles

Series & Parallel structures

How the arrangement of structures affect force and length change

Arrangement of fibres with muscle

Force-velocity relation, also power

For shortening & stretch

Springs connected in Series

Fixed position

Springs connected in Parallel

Fixed position

Fixed position

Fixed position

Structures in Series: Force at A and B are equal. For structures in series, forces do NOT add up

Force = A
Fixed position

Force = A+B
Fixed position

For structures in parallel, forces add up

Length changes
Length 2.0 m = 0.5 m Length 1.5 m

Connect in series
Length 4.0 m

= 1.0 m
Length 3.0 m

For structures in series, length changes add up

Length changes
Length 2.0 m = 0.5 m Length 1.5 m

Connect in parallel
Length 2.0 m

Length 1.5 m

= 0.5 m

For structures in parallel, length changes do NOT add up

Contractile and Elastic Structures

In series and in parallel

A muscle-tendon complex (MTC)

Bone

Parallel elastic component

Muscle fibres or Contractile Component (CC)

Tendon or Series elastic component (SEC)

Bone

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

Muscle or CC

Tendon or SEC

Bone

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 complicated connections can switch elasticity between series and parallel. Bone CC

PEC SEC Bone

Where and what are the SEC and PEC relative to the crossbridges?

Tendon (collagen) Aponeuroses (collagen) Epimysium (collagen) Filaments (titin) Filaments (myosin, actin)

series series parallel parallel series

Outline
Sliding filaments & the crossbridge cycle

Force & Power

Antagonistic muscles

Series & Parallel structures

How the arrangement of structures affect force and length change

Arrangement of fibres with muscle

Force-velocity relation, also power

For shortening & stretch

Arrangement of muscle fibres: some examples

parallel fusiform

triangular unipennate

bipennate multipennate

parallel

pennate

Arrangement of fibres within muscle:

Pennation increases muscle force

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 the pennate muscle as in the parallel muscle

Net effect: pennate muscle produces 2 * 0.87 = 1.74 times more force than the parallel muscle

Pennation reduces muscle shortening velocity

In each unit of time

the cos rule means that muscle shortening is cos * fibre shortening.

also each fibre in the pennate muscle only shortens half as far as each fibre in the parallel muscle.

Net effect: pennate muscle shortening is only 0.5 * 0.87 = 0.41 times as much as the parallel muscle per unit time

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

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

Before stimulation of the muscle

Muscle Force Muscle length (Lever movement) time

Isometric phase muscle force too small to lift weight

start stimulation of the muscle

Muscle Force
Muscle length (Lever movement) Stim time

Isotonic shortening: constant force during shortening

During stimulation, muscle force enough to lift weight

Muscle Force
Muscle length (Lever movement) Stim time

Larger weight
Before stimulation of the muscle

Muscle Force Muscle length (Lever movement) time

Isometric phase muscle force too small to lift weight

During stimulation of the muscle

Muscle Force
Muscle length (Lever movement) Stim time

Isotonic shortening: constant force during shortening

During stimulation of the muscle

Muscle Force
Muscle length (Lever movement) Stim time

Larger force & slower velocity

Inverse relation between force and velocity of shortening

The Force Velocity Curve

Power = work rate = (force x length ) / time = force x (length / time)

= force x velocity

0.2

1.0
Power

Force

0.5

0.1

0.0 0.0 0.5 1.0 Velocity 1.5

0.0 0.0 0.5 1.0 Velocity 1.5

Contraction with lengthening (eccentric)

The book is lowered in a slow, controlled movement. Biceps is acting as a brake. Biceps is producing force, EMG, etc, (=contracting) The elbow extends as the length of biceps increases due to the book & gravity. Work is done on biceps.

Force during isovelocity stretch of active Force-Velocity relation for Stretch muscle
stim

stim

stim

stretch
Velocity

shorten

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

Skeletal Muscle Basics

Contraction. Basic mechanical properties

Summary
Tension-length curve, max force at max filament overlap Cross bridge cycle, myosin head binds to actin, ATP splitting, repetitive

Muscle morphology:
short fat muscles, high force, low speed; long thin muscles, low force, high speed Inverse force-velocity relation Power = Force*velocity; max power at ca 1/3 max force or velocity Eccentric contractions, high force.

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.