Fatigue

study guide
Exercise Physiology:
Understanding the Athlete Within

Key concepts

Fatigue is a complex phenomenon involving interplay between central and

peripheral, and psychological and physiological, factors
Factors in fatigue include "central fatigue", substrate depletion, metabolite
accumulation, electrolyte disturbances, dehydration, hyperthermia and muscle
microtrauma
The factors that limit sprint and endurance performance differ markedly, resulting in
the adoption of different training and performance enhancement strategies
VO2 max and lactate threshold are used as measures of endurance performance
potential, with the latter perhaps the best predictor.

Fatigue
Fatigue can be defined as a "reduction in the force generating capacity" or "an inability
to maintain the required or expected force output". The latter is defined by specific task
failure, whereas the former is defined by a reduction in the maximal force generating
capacity prior to task failure.

Figure from D. G. Allen, G. D. Lamb, and H. Westerblad

Skeletal Muscle Fatigue: Cellular Mechanisms
Physiol. Rev. January 2008 88:287-332

Fatigue is a complex phenomenon that involves the interplay between central factors,
which are usually defined as those which affect the central nervous system, and those
peripheral factors which have a direct influence on skeletal muscle force and power
generation and the various physiological systems that support skeletal muscle.
There are complex interactions between central and peripheral factors (see figure right,
courtesy of Prof. T. Noakes & Dr. A. St. Clair Gibson, University of Cape Town, South
Africa) - feedback from contracting skeletal muscles appears to influence the central
motor drive to those muscles, thereby modifying exercise performance. If that feedback
is blocked, there is greater central drive and improved exercise performance initially, but
eventually premature fatigue due to local limitations within the contracting skeletal
muscles.

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Fatigue study guide

Exercise Physiology:
Understanding the Athlete Within

Potential factors involved in central nervous system fatigue include:

Neuroglucopenia (reduced glucose supply to brain)

Cerebral hypoxemia
Hyperthermia
Pain
Amphetamines, caffeine - modify sensations of fatigue
Experiences, emotions, motivation
Existence of "central governor"?

In terms of understanding the factors contributing to fatigue within contracting skeletal

muscle, review the basic steps in excitation contraction coupling, as these are potential
sites of fatigue. Some of the mechanisms causing fatigue here are summarized in the
figure below (courtesy D. G. Allen, G. D. Lamb, and H. Westerblad Skeletal Muscle
Fatigue: Cellular Mechanisms Physiol. Rev. January 2008 88:287-332).

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print or download this material solely for your own information, research or study.

Fatigue study guide

Exercise Physiology:
Understanding the Athlete Within
Much attention has focused on potential metabolic factors in fatigue - these involve
substrate depletion and/or metabolite accumulation, both of which impact on the rate of
ATP turnover. It is interesting to speculate on whether the decline in force is due to
reduced ATP turnover, or whether the rate of ATP turnover is reduced to a level entirely
appropriate for a force output that is lower due to some other fatiguing mechanism?

L. L. Spriet, K. Soderlund, M. Bergstrom, and E. Hultman. Anaerobic energy release in skeletal

muscle during electrical stimulation in men. J. Appl. Physiol. February 1, 1987 62:611-615

Strategies to enhance fatigue resistance include:

Training physical, technical, mental

Nutrition CHO, fluid, protein?
Heat acclimatization, cooling
Supplements, drugs? Gene doping?

Sprinting performance
Determinants of sprinting performance include:

Muscle mass to generate force and power

Fast twitch fibres
Neuromuscular recruitment
Fast reaction time
Ability to generate and tolerate acidosis (buffer capacity).

Factors contributing to fatigue during high intensity, sprinting exercise include:

Hyperkalemia increased ECF K+ due to release from contracting skeletal muscle

CP depletion dietary creatine supplementation is associated with increased CP
availability and improved sprint performance
Muscle glycogen depletion unlikely, although there may be loss of glycogen locally
at key sites of ATP utilization e.g. SR?
Acidosis (increased [H+]) some debate on direct effects of H+ on cross-bridge
cycling and force production, but does appear to reduce ability to maintain force
output which implies and effect on ATP turnover; induced alkalosis (increased ECF
pH) and increased muscle buffer capacity due either to sprint training or dietary
alanine supplementation (which increases muscle carnosine, a buffering molecule in
muscle) are associated with enhanced sprint exercise performance.

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print or download this material solely for your own information, research or study.

Fatigue study guide

Exercise Physiology:
Understanding the Athlete Within
Adaptations to sprint training include:

Increased muscle Na+/K+ ATPase and improved K+ regulation

Increased muscle buffer capacity
Enhanced muscle lactate/H+ transport capacity
Increased glycolytic enzymes
Increased VO2 max and muscle oxidative capacity
No good evidence of significant changes in muscle fibre type distribution.

Endurance performance
Determinants of endurance exercise performance include:

High VO2 max

Ability to maintain a high %VO2 max ("fractional utilization") during exercise
High power output at lactate threshold (LT) related to muscle oxidative capacity
Ability to oxidise power at high power outputs
Efficient/economical technique.

VO2 max sets the upper limit for aerobic energy production during exercise, whilst
muscle oxidative capacity is associated with the lactate threshold, fractional utilization
and the ability to maintain power output during prolonged exercise performance.

K. J. Davies, J. J. Maguire, G. A. Brooks, P. R. Dallman, & L. Packer

Muscle mitochondrial bioenergetics, oxygen supply, and work
capacity during dietary iron deficiency and repletion
Am. J. Physiol. Endocrinol. Metab. June 1, 1982 242:E418-E427.

The figure above summarises results from an experiment in which laboratory rats were
made severely iron deficient through dietary iron restriction. This reduces red cell mass
and [Hb] and muscle oxidative capacity. Iron was restored to the diet and rats recovered
over the next week. As you can see, VO2 max increased in line with the recovery in
haematocrit (%red cells in blood), consistent with the notion that oxygen delivery is the
primary determinant of VO2 max. Running endurance on the other hand, was restored in
parallel with the recovery in muscle oxidative capacity (as measured by muscle pyruvate
oxidase).
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print or download this material solely for your own information, research or study.

Fatigue study guide

Exercise Physiology:
Understanding the Athlete Within
There is a high correlation between muscle oxidative capacity and the lactate threshold
(LT). For this reason, the measurement of blood lactate levels during submaximal
exercise, including LT determination, is commonly used in the routine assessment of
endurance athletes. There is strong, positive correlation between exercise time to
fatigue and LT. This association may be partly related to the inverse relationship
between muscle glycogen use during exercise and LT.
Potential factors in fatigue during endurance exercise include:

"Central fatigue"
Hyperkalemia and loss of muscle potassium (although the increase is smaller in
magnitude with each contraction, over time the cumulative effect can be significant)
Reduced SR Ca2+ release lower ATP and glycogen close to the SR may affect
Ca2+ release and uptake by and from the SR
Muscle glycogen depletion
Hypoglycemia which results in reduced muscle CHO oxidation and neuroglucopenia
Dehydration impaired cardiovascular and metabolic function
Hyperthermia.

Carbohydrate and fluid ingestion enhance endurance exercise performance via

mechanisms that appear to be independent and additive.
Adaptations to endurance exercise training with implications for performance:

Increased VO2 max increased maximal stroke volume an cardiac output

Expanded blood volume
Increased muscle capillary density
Increased muscle oxidative capacity
Increased muscle Na+/K+ ATPase and improved K+ regulation
Reduced reliance on muscle glycogen and blood glucose and the same energy
expenditure; increased fat oxidation
Possible increase in type I fibres?

If you wish to read more on the physiology of endurance exercise performance, the
following article may be of interest:

Joyner, M.J. and E.F. Coyle. Endurance exercise performance: the physiology of
champions J. Physiol. 586: 35-44, 2008 http://jp.physoc.org/content/586/1/35.long

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print or download this material solely for your own information, research or study.

Fatigue study guide

Exercise Physiology:
Understanding the Athlete Within

Abbreviations
A

arterial

ADP

adenosine diphosphate

AP

action potential

ATP

adenosine triphosphate

CHO

carbohydrate

CON

control/placebo trial

CP/PCr

creatine phosphate/phosphocreatine

ECF

extracellular fluid

FV

femoral vein

GLY

glycogen

Hb

haemoglobin

HR

heart rate

LT

lactate threshold

Pi

inorganic phosphate

QO2

muscle oxidative capacity

ROS

reactive oxygen species

RyR

ryanodine receptor or SR Ca2+ release channel

SM

skeletal muscle

SR

sarcoplasmic reticulum

TT

transverse tubule

VO2

oxygen uptake

VO2 max maximal oxygen uptake

Image credits
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Man - Exhaustion by Red Rose Exile CC BY-SA 2.0 via Flickr

Diagram of the brain 1 - Figure courtesy of Prof. T. Noakes & Dr. A. St. Clair Gibson, University of
Cape Town, South Africa
Diagram of the brain 2 - Figure courtesy D. G. Allen, G. D. Lamb, and H. Westerblad Skeletal
Muscle Fatigue: Cellular Mechanisms Physiol. Rev. January 2008 88:287-332
Pair of graphs- x-axis= contraction time, sec - L. L. Spriet, K. Soderlund, M. Bergstrom, and E.
Hultman. Anaerobic energy release in skeletal muscle during electrical stimulation in men. J. Appl.
Physiol. February 1, 1987 62:611-615
Graph- recovery vs dietary iron repletion - Figure courtesy D. G. Allen, G. D. Lamb, and H.
Westerblad Skeletal Muscle Fatigue: Cellular Mechanisms Physiol. Rev. January 2008 88:287-332

Unless otherwise indicated, this material is The University of Melbourne. You may save,
print or download this material solely for your own information, research or study.