Muscle Physiology & Dynamics of Work

How a Working Horse Works

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Equine Muscle Physiology & Mechanics

Muscle Tissue Intro

Structure & Function
ƒ Muscle Microanatomy & Physiology
ƒ Dynamics of Work
ƒ Specific Muscle Fibers & Energy Substrates
Exercise & Effects on Muscle

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Muscle Tissue: Introduction

Muscles = Contraction

ƒ 3 Types of Muscle
• Visceral Muscles (Smooth Muscle) Involuntary
¾ GI Tract, Blood Vessels, Uterus, etc.
• Cardiac Muscle Involuntary
¾ Heart
• Skeletal Muscle (Striated Muscle) Voluntary
¾ Movement of Joints, Limbs, etc.
– Explosive power
– Stamina
– Motor Control

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Skeletal Muscle: Structure & Function

ƒ Large part of body weight (up to 40% including H20)

ƒ Closely associated with the skeletal, nervous, and
circulatory systems
• Manipulation impacts a range of tissues & systems
ƒ Generates heat
ƒ Each muscle is a collection of fibers & associated tissues
ƒ Attached to bone via tendons & connective tissue
• Least moveable attachment = origin
• Most moveable attachment = insertion

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Microanatomy
& Physiology

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Muscle Fiber = Individual Muscle Cell
ƒ Multinucleated – composed of fused cells
ƒ Large cells
• 10 – 100 µm diameter
• Approx 20 cm in length

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Muscle Cells

ƒ Specialized to contract
• Generate FORCE and MOVEMENT
ƒ Do not divide
• Increased muscle size is due to Increased cell size

Key Qualities of Muscle Cells

• Excitable
• Conductive
• Contractile

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Muscle Cell Key Components
ƒ Membrane = Sarcolemma
ƒ T-Tubules
• Transmit Messages
ƒ Mitochondria
• Generate Energy
• Numerous
ƒ Myofibrils
• 2 Proteins in long strands
• Heart of the contractile function
ƒ Sarcoplasmic Reticulum (Endoplasmic Reticulum)

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Muscle Cell & Associated Structures

As visible with a standard light microscope

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Skeletal Muscle

Electron Micrograph

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Dynamics of Work

¾ Mechanism of Contraction
¾ Stimulus of Contraction
¾ Energy for Contraction

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Sarcomere = Smallest Unit of Contraction

ƒ Repeating Pattern of Striations

ƒ Thick and Thin Filaments
Actin (Thin) & Myosin (Thick)
ƒ Myofilaments arranged in a specific pattern
H-Zone
Z-Line
A-Band

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Actin & Myosin

ƒ 2 Principal Muscle Proteins

ƒ Found in Myofibrils
ƒ Arranged in a Ring-like Structure
• Usally 6 Actin strands around a Myosin fibril
ƒ Run Parallel & Lengthwise
ƒ Myosin (Thick) has protrusions (Crossbridges)
ƒ Actin (Thin) is intertwined with thinner topomyosin and
troponin

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Mechanism of Contraction
1. Nerve Impulse Stimulation
2. CA++ Released into Cytoplasm by Sarcoplasmic
Reticulum
3. CA++ Binds to Troponin, which Rotates
4. Tropomyosin Moves and Actin is Exposed to Myosin
5. Myosin Crossbridge Binds to Actin
6. Crossbridge Drags Along Actin (Power Stroke)

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When all the crossbridges in a sarcomere act together,
the whole sarcomere contracts

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Mechanism of Relaxation

7. Nerve Impulse Ends

8. SR Reabsorbs CA++
9. CA++ Dissociates from Troponin
10. ATP Binds to the Crossbridge
11. Crossbridge Disconnects from Actin
12. Actin Fibers Return to Previous Positions
13. Sarcomere Relaxes

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Contraction-Relaxation
ƒ A muscle cell may not go back to immediate complete
relaxation
ƒ Contraction can continue through a series of stimulations
(Summation)
ƒ Summation increases the total force of contraction
ƒ If the stimulus is great enough, many sarcomeres in
many fibers are recruited, and the muscle as a whole
contracts.
ƒ Allows for varying amounts of work
ƒ Muscle failure occurs when the maximum number of
fibers are stressed beyond their limits

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Stimulus of Contraction:
Muscle Contraction is Controlled by Motor Nerves

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Interaction of Motor Nerves and Muscle Fibers

ƒ Each muscle is innervated by only one motor nerve

ƒ One nerve can innervate a number of muscles
ƒ Each nerve controls many fibers (motor units), the fewer
the fibers the more delicate the movement
ƒ If nerve contact is lost, fibers shrink (atrophy)
ƒ The pattern of nerve activity determines the fiber type

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Feedback Loop
ƒ Feedback from the tendon and stretch receptors
controls motor nerve activity
ƒ Motor nerve activity is also controlled by higher
centers (brain)

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Mechanism of Neuromuscular
Stimulation (Excitation)
1. Electrical depolarization occurs along the stimulated
nerve
2. Nerve end touching the muscle fiber releases a
neurotransmitter (ACH)
3. Depolarization of the muscle cell membrane (Action
Potential)
4. T-tubules open in SR and Ca++ is released
5. Increased intracellular Ca++ allows actin and myosin to
interact and the crossbridge cycle starts

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Relaxation

ƒ When electrical activity stops, the calcium is removed

and contraction stops
ƒ Muscle must relax between each contraction by actively
pumping Ca back to SR
ƒ Ion pumps in the cell membrane actively repolarize the
muscle cell membranes
ƒ All processes necessary for relaxation are active –
require energy

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Energy for Contraction
ƒ Each crossbridge requires ATP
ƒ Each myosin strand has dozens of crossbridges
ƒ Each muscle fiber has hundreds of myosin strands

¾ Muscle Contraction Requires Significant Energy

Basic Unit of Energy = ATP

ATP Æ ADP & Pi ÖENERGY

(ATP + H2O Æ ADP + Pi +H+ + Energy)
ATP= adenosine triphosphate; ADP=adenosine diphosphate;
Pi=Inorganic phosphate
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For a horse to maintain exercise for more than a few
seconds, ATP stores in muscle must be replenished at
an appropriate rate.

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Energy for Contraction

Fuels
ƒ Intramuscular Triglycerides & Glycogen
ƒ Extracellular FFAs from Adipose Deposits and Glucose from the
Liver

Total amount of fuel stored in a 1,000 lb horse

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Two Main Pathways For Energy Metabolism

ANAEROBIC

AEROBIC

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Aerobic Metabolism

ƒ Occurs in Mitochondria
ƒ For low energy demands of slow speed exercise
ƒ Primary pathway for endurance exercise
ƒ Gallop speeds < 18sec/200m can usually be met by
aerobic metabolism in fit horses

ƒ Training can increase capacity to generate energy

aerobically
• Enhanced oxygen delivery to muscle
• Increased mitochondrial density
• Increased enzyme concentrations

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Aerobic Metabolism
ƒ Oxidative Phosphorylation
ƒ Fats & CHO oxidized to produce ATP
Fats – stored in depots around body
CHO – stored as glycogen in liver & muscle
(glycogen metabolizes to glucose)
Aerobically metabolized approx 2x as fast as fat

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Aerobic Metabolism

Limitations
ƒ Primarily limited by availability of oxygen in working
muscles
ƒ Upper airway obstructions
ƒ Cardiovascular system impairment
ƒ Hemoglobin concentration

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Anaerobic Metabolism

ƒ High intensity exercise of short duration (2-3 min in

horses)
ƒ At start of fast exercise, O2 delivery does not
immediately reach the level required to support aerobic
metabolism
• Approx 30-45 seconds of exercise is required before maximal
rate of oxygen use is achieved
• During this time, anaerobic metabolism supplies energy
ƒ Fast Exercise not totally anaerobic; makes up the deficit
• Horse galloping over 1200 m, aerobic metabolism provides
approximately 70% of energy

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Anaerobic Metabolism
ƒ Glycolysis = Degradation of muscle glycogen
to lactate
ƒ Results in increases in lactate, hydrogen ions
and Pi in the cells
ƒ Lactic acid accumulation and fatigue develop
as muscle pH falls
¾ At pH < 6.4 glycolysis and contraction are
inhibited

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Different Muscles have Fibers with
Different Properties
Type I & Type IIA
ƒ High Oxidative Capacity
ƒ Store Triglycerides & Glycogen
ƒ Standing and posture: Slow contracting fibers that are well supplied
with oxygen – example stay apparatus
ƒ Type I aka “Slow Twitch” Fibers “Red Fibers”

Type IIB
ƒ Low Aerobic Capacity
ƒ Store Glycogen
ƒ Athletic Movements: Muscles that generate rapid movement contain
fast fibers and can work for short periods without oxygen
ƒ Type II aka “White” Fibers, “Fast Twitch” Fibers

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Fiber Type Recruitment Based on Energy
Requirements

Walking
ƒ Primarily Type I Fibers
• primarily aerobic energy, primary substrate is fat

Transition from Walk to Trot and Cancer

ƒ Type IIA Fibers Recruited
ƒ Primarily aerobic energy, substrate is both fat and glycogen

Transition to Gallop
ƒ Type IIB Fibers Recruited
ƒ Energy no longer purely aerobic,

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Exercise

ƒ Concentric Exercise
• Isometric – constant length
• Isotonic – constant force
• Or a mixture of the two

ƒ Eccentric Exercise
• Lengthening contractions

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Effects of Exercise on Muscle

ƒ Lack of exercise leads to fiber atrophy

ƒ Gentle exercise maintains muscle mass & flexibility
ƒ Moderate long term activity increases fatigue resistance
ƒ High load exercise leads to muscle fiber hypertrophy

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Muscle Fatigue

ƒ Prolonged and/or strong contraction > Fatigue

• Inability of contractile and metabolic processes to
continue supplying the same work output
ƒ Nerve sends electric stimulation, NMJ transmits, action
potentials spread over muscle fibers
¾ However contraction becomes progressively weaker due
to reduced ATP in the muscle fibers
¾ Interruption of blood flow through a contracting muscle
leads to almost complete fatigue in less than a minute
due to loss of nutrient supply

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Muscle Fatigue

Endurance Horses
ƒ Most often due to glycogen depletion, as most work is
performed aerobically

Race Horses
ƒ Most often due to lactic acid accumulation

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Lactic Acid or Lactate

ƒ By product of anaerobic glycolysis

ƒ A potential cause of late onset muscle soreness 24 – 48
hours after intense exercise
ƒ Sent from muscle to blood and removed via liver
ƒ Removal requires oxygen and is hastened by light work
during recovery

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Muscle Atrophy

ƒ Results anytime a muscle is not used or used only for

weak contractions
ƒ Denervated muscle begins immediate atrophy
• Example: Sweeney
¾ Injury to Suprascapular N causing atrophy in supraspinatus &
infraspinatus

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Muscle Hypertrophy

ƒ Diameter of individual muscle fibers increase

ƒ Sarcoplasm increases
ƒ Fibers gain in nutrient and intermediary metabolic
substances (ATP, creatine phosphate, glycogen,
intracellular lipids, additional mitochondria)
ƒ Myofibrils may also increase in size
¾ Hypertrophy increases both power of the muscle and the
nutrient mechanisms to maintain that power

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Conclusion

ƒ Muscle Microanatomy & Physiology

ƒ Dynamics of Work
ƒ Specific Muscle Fibers & Energy
Substrates
Together IMPACT > Exercise &
Its Effects on Muscle

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