Lactate measurements in equine sport science
Lactate measurements in equine sport science
Accumulation
of lactate in muscle cells and in the blood is a normal consequence of fast exercise in
the horse. At low speeds, the horse is able to generate sufficient energy by
catabolism of glycogen, glucose and fat. This metabolic process uses oxygen to
generate ATP, and is referred to as aerobic. At higher speeds, aerobic
metabolism does not regenerate ATP quickly enough. Pyruvate accumulates in
muscle cells, and it is converted to lactate ions at increased
rates.
Exercise
at speeds greater than approximately 700–800 m/min recruits fast twitch skeletal
muscle fibers. These fibers can be classified as being highly oxidative or
highly glycolytic in nature. At fast speeds, recruitment of fast twitch, highly
glycolytic muscle fibers results in accumulation of lactate anions and hydrogen
ions in the muscle cells, due to the contribution of anaerobic glycolysis to ATP
resynthesis. Both these ions diffuse into the extracellular fluid. It is
generally thought that the stimulus for anaerobic glycolysis in skeletal muscle
fibers during fast exercise is a limitation to the supply of oxygen at the
cellular level.
The
production of many molecules of lactate and hydrogen ions results in acidosis of
both the skeletal muscle cells and the blood. The increasing acidity of muscle
cells is implicated in fatigue during intense exercise, but the direct cause and
effect is still debated. In any horse at top speed for about 800 m, the
accumulation of lactate and the concomitant cellular acidosis has a negative
effect on energy production by anaerobic glycolysis. The rate of ATP production
is decreased, and the animal reduces speed.
Resting
blood lactate concentration in the horse is approximately 1–1.5 mmol/L. At low
speeds this value does not change greatly from the resting value. At moderate
speeds lactate begins to accumulate in the blood. Accumulation of lactate in
blood occurs most quickly when the work speed is faster than that at which blood
lactate is about 4 mmol/L. This work speed at which blood lactate is 4 mmol/L is
often referred to as the anaerobic threshold, or the speed at onset of
accumulation of blood lactate (OBLA). It is also frequently referred to
as VLa4. VLa4 is therefore the work velocity
which results in a blood lactate of approximately 4 mmol/L. This value is
derived from inspection of graphs of exercise speed (on the X axis)
plotted against blood lactate concentration (on the Y axis). The rate of
lactate accumulation generally parallels the accumulation of adrenaline in the
blood.
At
speeds greater than VLa4, lactate accumulates rapidly in the
blood. The general relationship between velocity and blood lactate is therefore
usually described as exponential. However, if sufficient steps are used in the
exercise test, the relationship is described by two straight lines, with an
obvious velocity at which the blood lactate begins to accumulate in
blood.
After
a race, blood lactate concentrations are usually greater than 20 mmol/L. It is
normal for the blood and muscle lactate concentration to then gradually decrease
over a 1–2 h period after a race or fast work. Acidosis of muscle and blood is a
normal result of fast work, and this acidosis is rapidly reversed by the horse’s
own metabolism. Racehorses do not develop chronic acidosis due to lactate
accumulation in training, and therefore alkaline supplements are not
required.
The
rate of decrease in blood lactate after exercise is affected by the activity
during this period. For example, blood lactate decreases more quickly after
strenuous exercise if the horse is trotted for 3 0minutes, rather than walked.
However, such a practice might delay cooling of the horse.
Many
studies of horses trained on both treadmills and on racetracks consistently
demonstrate that training results in lower blood lactate concentrations at the
same work speed. The speed at which blood lactate begins to accumulate
rapidly, VLa4, also increases. The horse is able to work at a
higher speed without accumulating lactate.
Repeated
tests of the blood lactate relationship with velocity are suitable as a means of
measuring increasing stamina with training. If more than one horse is
tested, measurements of VLa4 enable comparisons of the
relative stamina in each horse. VLa4 measurements every 2–3
weeks also enable measurement of changes
in fitness through the training program.
However, VLa4 measurements have the disadvantage of requiring
several blood collections and analyses. A simple, one step exercise test can be
designed to give the same information. For example, the blood lactate
concentration 3 minutes after a two minute gallop on a treadmill was correlated
with racing performance in thoroughbreds (Evans et al., 1993). The exercise test
for race-fit horses was as follows. The treadmill angle was set at 6° (or a 1 in
10 slope). The horse is trotted for 2 min (at 4 m/s), followed immediately by 2
min slow cantering (6 m/s). The horse is then walked on the treadmill for 4 min,
and then given 2 min exercise at 10 m/s. Racehorses with superior stamina, or
endurance fitness, have blood lactate concentrations of <4 mmol/L after the
test.
The
blood lactate response is therefore a guide to racing ability, and the other
factors which contribute to racing success must not be ignored. However, a
treadmill study of 12 English Thoroughbred racehorses indicated that 47% of the
variability in Timeform rating (a handicap rating system) was due to
variability in their blood lactate response to treadmill exercise, using a test
similar to that described above. The better horses had a lower blood lactate
after treadmill exercise at 10 m/s on a treadmill inclined at 10%. Similar
results have been reported in trotting horses. The blood lactate response to
exercise is a very important determinant of likely success or failure on the
racetrack.
When
blood is collected for lactate assays, it should be added to tubes containing a
suitable anticoagulant and inhibitor of glycolysis. Fluoride and oxalate
combinations are suitable, as the blood stores well at room temperature for at
least two days in those chemicals. However, if possible, blood should be stored
in a refrigerator or on ice until analysis. Individual laboratories should be
consulted concerning ideal storage conditions, as different centres use
different techniques to conduct the analysis.
Lactate
assays can also be conducted on plasma. However, plasma and whole blood lactate
concentrations in blood collected after exercise are not equivalent. Plasma
lactate concentrations are 140–150% of concentrations found in whole blood, due
to unequal distribution of lactate between plasma and erythrocytes in horse
blood after exercise rainger etal., 1995). Plasma and whole blood lactate
concentrations should therefore not be directly compared.
In
the past, lactate assays have been complicated and not readily available. The
analysis of plasma or serum for lactate concentration has been greatly
simplified by the development of rapid analysers. However, these technologies
should be used with great caution. They express a value for blood lactate, but
blood lactate is not directly assayed. The assay is of plasma lactate, and if
haematocrit exceeds 50% large errors occur. (Evans and Golland 1996). After
exercise haematocrit will usually exceed 50% in horses.
Incremental
treadmill exercise tests, with measurements of heart rate and blood lactate at
various speeds, can also be conducted. Such tests enable calculation of indices
of fitness such as V200 and VLa4. Collection
of blood at each speed necessitates stopping the treadmill after each step, or
placement of an IV catheter. Sterile heparinized saline solutions can be used to
maintain catheter patency during the tests. Such tests are now common at
research centres with treadmills for assessment of poor performance.
Measurements of heart rate, blood lactate and oxygen consumption are usually
conducted during submaximal exercise and during exercise at maximal intensities.
However, blood lactate concentrations after maximal exercise have not been
correlated with racing performance.
Blood
lactate measurements after field exercise are not advisable for assessments of
fitness in thoroughbreds. Small variations in speed or distance of the gallop
can have large effects on the post-exercise blood lactate
concentration.
Blood
lactates after exercise have been recommended for guiding the training
intensity. A recent study found no difference to the aerobic fitness produced
after training at lactate concentations of 2 or 4 mmol/L. However, post exercise
blood lactates may be useful for guiding training at higher intensities. It is
known that muscle adaptations to training occur at 80-100% of HRmax. At these
intensities PLASMA lactate concentrations immediately after exercise could be
expected to be in the range 10-15 mmol/L.
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