MUSCLE FATIGUE IN THE HORSE: A REVIEW
Melyni Worth, MSc
SUMMARY
The onset of fatigue within a muscle depends on several
factors, including the duration or intensity of work and
amount of training. In aerobic work over prolonged time
it appears to be the level of glycogen stored within the
muscle fibers that becomes limiting. In anaerobic work
the limiting factor may be a combination of the amount
of substrate available to the cell and the ability of the cell
to continue working at low pH. Further work is needed to
establish which is the most limiting of these two factors,
levels of stored glycogen or accumulating levels of lactate
and concomitant lowering of muscle pH.
LITERATURE REVIEW
With the rise in popularity of equine athletic
competitive events, there has been an associated
increased interest in optimizing training methods, and in
defining the factors which limit equine performance.
There are many factors, both mental and physical, which
affect the performance of an athlete. These factors
include level of skill, confidence of fitness and mental
well-being, but the ultimate limit to all physical effort is
the onset of fatigue. Morehouse and Miller ~2 defined
three kinds of fatigue. The first type, chronic, is due to
mental stress, boredom or anxiety. The second, nervous
or central, could be due to an increase in stimulus
threshold of neurons, depletion of Ca or K ions or as
postulated by Asmussen,~ an inhibition elicited by
nervous impulses from receptors on fatigued muscles.
This inhibition is thought to arise from reticular
formation and can be overridden by other signals of
Author's address: Department of Animal Science, Virginia
Polytechnic Institute and State University, Blaeksburg, VA 24061.
276
peripheral or central origin, which may explain the role
of motivation, whips, or certain stimulant drugs on
extending the onset of fatigue past the normal point. The
third type of fatigue is muscular or peripheral fatigue
which is defined as the point at which the muscle fiber will
no longer respond to nervous stimulation. This inability
to respond may be due to several factors, such as
depletion of energy substrates and accumulation of
metabolic intermediates. Accumulation of metabolites
may have inhibitory effects on enzyme activities and
produce an increase in stimulatory threshold due to
accumulation of K or Ca ions on the extra-cellular and
extra-reticular compartments.
Muscle contraction is brought about by the sliding of
actin and myosin filaments over one another; the energy
for this contraction comes from the dephosphorylation
of ATP - A D P + Pi. The ADP is then rephosphorylated,
initially via creatine kinase which catalyzes the reaction
phosphocreatine + A D P - creatine + ATP. It has been
postulated that the creatine stores in the equine will
provide A T P regeneration for the first 10 seconds of
exercise. After that, the ADP must be rephosphorylated
via either oxidative phosphorylation (aerobic) or
glycolysis (anaerobic).
Oxidative p h o s p h o r y l a t i o n occurs within the
mitochondria and utilizes acetyl CoA, which is derived
from either glycolysis or B-oxidation of fats. The
ultimate electron acceptor is oxygen. There are 36 ATP's
generated for each molecule of glucose oxidized via
aerobic metabolism. This is the major pathway for energy
production in prolonged work at slower paces. This
highly efficient oxidative phosphorylation occurs in all
types of exercise provided there is sufficient oxygen
available.
Glycoloysis, which is the oxidation of glucose to
pyruvate and subsequently to lactate, yields only 2 ATP's
EQUINE VETERINARY SCIENCE
per molecule of glucose, but is independant of oxygen
supply and therefore provides energy under conditions of
limited oxygen availability. The rephosphorylation of
ATP via glyeolysis is the major pathway of energy supply
in animals performing fast, explosive work such as
sprinting, jumping or fast running.
Most recent research in exercise physiology of horses
has examined the effect of aerobic work on endurance
horses, but some workers are currently studying the
effects of more intense anaerobic work patterns.
Lindholm and Piehl 6 investigated muscle structure and
the electrolyte and metabolite concentrations in muscles
of Standardbred horses at varying ages and states of
training. They reported three types of muscle fibers: Type
I or slow twitch (ST) which have a high oxidative
capacity and use fat or glucose as a fuel substrate and
Type II or fast twitch fibers. The Type II fibers were
further divided into Type IIA which have an
intermediate-to-high oxidative capacity (FTH) and Type
l i b which have a low oxidative capacity (FT). They also
reported that the horse has a very high level of glycogen
stored within the muscle compared to man or dog.
Muscle glycogen levels are also found to be closely
correlated with age and level of training. The various
fiber types and their relative oxidative and fuel capacities
have been further studied by several groups of workers.23
15 16 17 20
Physical exertion affects various physiological
functions. Biochemical alterations which are reflected in
changes in plasma constituents are easiest to assess. Most
early studies were conducted using privately owned
horses which were competing in various events. 4 s 9 to ~3~4
is t9 All workers found that with long-duration aerobic
exercise there was a decrease in levels of plasma glucose,
and an increase in total proteins, lactate, free fatty acids
(FFA) and hematocrit. Depending on the speed and
environmental temperature, there were shifts in levels of
K, Na, Mg and Ca, but these varied considerably among
studies. Problems included the considerable variation in
the types of horses, level of training and fitness, feeding
regimen, and the inability to do follow-up work due to
private ownership.
Snow and coworkers 2 conditioned a group of
Thoroughbred horses through a course of endurance
training. These horses then competed in a mock
endurance-type event. It was reported that the parameter
which best correlated with the degree of fatigue was
depletion of muscle glycogen. Hodgson et alp studied
muscle glycogen depletion and repletion patterns of
horses competing in endurance events of varying
distances. They reported that the distance completed was
highly correlated with the type of muscle fiber recruited
and the level of glycogen present in those fibers. Both
groups of workers reported that in endurance work the
Type I (ST) fibers are recruited first, followed by Type
IIA (FTH) and finally Type l i b (FT). In aerobic exercise,
only when the first two types of fibers are depleted of
glycogen is the last type brought into use.
Lindholm 7 studied progressive recruitment of fibers at
increasing speeds of exercise in Standardbred horses. He
concluded that at lower speeds of work the Type I fibers
were preferentially recruited but, as speed of exercise
Volume 5, Number 5
increased, so did the level of recruitment of Type IIA and
B fibers. In all cases, the point at which a fiber was no
longer used was when it was depleted of glycogen. In
resting animals the Type I fibers were lower in glycogen
than Type II fibers, but during long exercise at slow
speeds Type I fibers were preferentially recruited,
indicating their ability to utilize F F A as a fuel substrate.
However once the fiber was depleted of glycogen it could
no longer function despite adequate levels of circulating
FFA.
During short term intense exercise, e.g., galloping at
maximum pace, greater than or equal to 100 percent VO2
max, anaerobic sources of A T P regeneration were the
most important. As a result there was an accumulation of
lactate, both within the muscle cell and in blood serum
due to diffusion. At these work loads, the rate of diffusion
out is less that the rate of formation and as a result there is
marked accumulation within the cell. In association with
the lactate accumulation there is a buildup of hydrogen
ions which results in a decrease in intracellular pH. At
least two enzymes are inhibited by low pH,
phosphofructokinase (PFK) and myosin ATPase, thus
impairing muscle functioning.
In man high levels of muscle lactate are associated with
pain. H Although pain is difficult to measure in horses,
performance may depend partially on the animals
perception of and willingness to endure pain. The
importance of muscle p H has been described by Sutton et
al. 2~ Sodium bicarbonate was administered to subjects
shortly before undergoing intense exercise. The
administration of bicarbonate resulted in a very slight
alkalosis and a prolonged time to fatigue, although
similar levels of lactate were present in both treated and
controls. Sutton and coworkers suggested that the
bicarbonate was also reaching higher than normal
concentration within the muscle fibers and increasing
their buffering capacity.
REFERENCES
h Asmussen E: Muscle fatigue. Med Sci Sports 11:313, 1979.
2. Essen B, Lindholm A, Thorton J: Histochemical properties of
muscle fiber types and enzyme activities in skeletal muscles of
Standardbred trotters. Equine Vet J t2:175, 1980.
3. Gunn H M: Differences in the histo-chemical properties of skeletal
muscle of different breeds of horses and dogs. J Anat 127:615, 1978.
4. Grosskopf JFW and Van Vensburg J J: Some observations on the
haematology and blood biochemistry of horses competing in 80 km
endurance rides. In Equine Exercise Physiology Eds DH Snow, R Rose
and S Persson. Granta Editions, Cambridge, 1983.
5. Hodgson DR, Rose R J, Allen JR: Muscle glycogen depletion and
repletion patterns on horses performing various distances of endurance
exercise. In Equine Exercise Physiology Eds DH Snow, R Rose,and S
Persson, Granta Editions, Cambridge, 1983.
6. Lindholm A, Peihl K: Fibre composition enzyme activities and
concentrations of metabolites and electrolytes in muscles of
Standardbred horses. Act Vet Scand 15:287, 1974.
7. Lindholm A: Metabolic response and muscle metabolites during
different exercise intensities in trotting horses. Acta Physiol Scand
Suppl. 396, 1979.
8. Lucke JN, Hall GM: Biochemical changes in horses during a 50
mile endurance ride. Vet Rec 102:356, 1978.
9. Lucke JN, Hall GM: Long distance exercise in the horse: Golden
Horseshoe Ride 1978. Vet Rec 106:405, 1980.
10. Lucke JN, Hall GM: Further studies on the metabolic effects of
long distance riding. Golden Horseshoe Ride 1979. Equine Vet J
123:189, 1980.
277
 11. Osnes JB, Hernansen L: Acid-base balance after maximal
exercise of short duration. J Appl Physiol 32:59, 1972.
12. Morehouse LE, Miller Jr AT: Physiology o f Exercise CV Moseby
Co, St. Louis, 1976.
13. Rose RJ, Purdue RA, Hensley W: Plasma biochemistry
alterations in horses during endurance riding. Equine Vet J9" 122, 1977.
14. Rose R J: Haematological changes associated with endurance
exercise. Pet Rec 110:175, 1982.
15. Snow DH, Guy PS: Percutaneous needle muscle biopsy in the
horse. Equine Vet J 8:150, 1976.
16. Snow DH, Guy PS: Muscle fiber type composition of a number of
limb muscles in different types of horse. Res Vet Sci 28:137, 1980.
17. Snow DH, Guy PS: Fibre type and enzyme activities of the
gluteus medius in different breeds of horse. In Biochemistry o f Exercise
IV-B Eds J Poortmans and G Niset. University Park Press, Baltimore.
pp. 275-282, 1981.
18. Snow DH, Makensie G" Some metabolic effects of maximal.
exercise in the horse and adaptations with training. Equine Vet J 9:134,
1977.
19. Snow DH, Rose R: Hormonal changes associated with long
distance exercise. Equine Vet J 13:195, 1981.
20. Snow DH, Kerr MG, Nimmo MA, Abbott EM: Alterations in
blood, sweat, urine, and muscle composition during prolonged exercise
in the horse. Vet Rec 110:377, 1982.
21. Sutton JR, Jones NL, Toews OJ: Effect of pH on muscle
glycolysis during exercise. Clin Sci 61:331, 1981.

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