Published December 4, 2014
Maximal lactate steady state during exercise in blood of horses1
A. E. Lindner2
Arbeitsgruppe Pferd, c/o Heinrich-roettgen-Str. 20, D-52428 Juelich, Germany
ABSTRACT: The speed producing the maximal lactate steady state (maxLASS) is supposed to be the optimal speed to condition for endurance. The maxLASS
was defined as the maximal speed at which the blood
lactate concentration ([LA]) between the 5th and the
25th min of continuous exercise did not increase by
more than 1 mmol/L. According to the aerobic-anaerobic lactate threshold concept determined in humans,
maxLASS corresponds to v4 [speed in a standardized
exercise test (SET) shown to produce an [LA] of 4
mmol/L; generalized to vi for the speed producing an
[LA] of i mmol/L]. Four Thoroughbreds were submitted
to a treadmill-based SET to determine their blood lactate-running speed (BLRS) relationship and calculate
the individual v1.5, v2, v2.5, v3, and v4 values (velocities
run under defined conditions inducing 1.5, 2, 2.5, 3, and
4 mmol/L of blood LA). Afterward, horses ran on the
treadmill for 40 min at their v1.5, v2, and v2.5 every 3 d.
Another 14 horses were submitted to SET in the field
to determine their BLRS relationships and to calculate
their v2. The day after the SET, these horses ran once
between 15 and 30 min at their v2. In the horses that
ran on the treadmill, maxLASS only occurred when
running at their v1.5. Blood [LA] did not increase by
more than 1 mmol/L between the 10th min and the
end of exercise for all the horses that ran in the field at
their v2. These data indicate that maxLASS of horses is
not greater than v2 and therefore less than in running
humans.
Key words: blood, exercise, horse, lactate, maximal, steady state
©2010 American Society of Animal Science. All rights reserved.
INTRODUCTION
For more than 30 yr, points on the blood lactate vs.
speed or power curves have been used in humans to define the transition from aerobic to partly anaerobic energy metabolism, as indicators of aerobic performance
and as a means of predicting exercise intensities for endurance training (Mader et al., 1976; Keul et al., 1979;
Sjödin and Jacobs, 1981; Stegmann and Kindermann,
1982; Simon, 1986; Tegtbur et al., 1989). In the 1980s,
the v4 [speed in a standardized exercise test (SET)
shown to produce a blood lactate concentration ([LA])
1
The author is very grateful to the Verein zur Förderung der
Forschung im Pferdesport (Jülich, Germany), the Wissenschaftliche Gesellschaft der Schwarzwald-Tierklinik (Jülich, Germany),
the Höveler Kraftfutterwerke GmbH (Jülich, Germany), and the
HODIBE Reitsport GmbH (Jülich, Germany) for the material and
financial support; to the Institut für Klinische Biochemie of the University of Bonn for conducting the lactate analysis; and to all the
members of the Arbeitsgruppe Pferd who helped to complete this
study, especially Jutta Werkmann, Matthias Sobotta, and Peter von
Wittke. Gratitude is also extended to Kenneth McKeever for revising the manuscript.
2
Corresponding author: [email protected]
Received November 24, 2009.
Accepted February 23, 2010.
J. Anim. Sci. 2010. 88:2038–2044
doi:10.2527/jas.2009-2693
of 4 mmol/L; generalized to vi for the speed producing
an [LA] of i mmol/L] threshold suggested by Mader et
al. (1976) was adopted for use in the horse (Isler et al.,
1982; Persson, 1983; Wilson et al., 1983; Straub et al.,
1984; Snow, 1987).
The aerobic-anaerobic lactate threshold is supposed
to identify the maximal intensity of exercise at which
blood lactate production and clearance during exercise
is at a balance, known as maxLASS. The intensity
to achieve maxLASS is believed to be optimal to improve endurance (Mader et al., 1976). However, this
assumption has not been demonstrated yet (Heck and
Beneke, 2008). Heck et al. (1985) showed in running
humans that the v4 corresponded to maxLASS. They
defined maxLASS as the maximal speed at which the
[LA] does not change by more than 1 mmol/L between
the 5th and the 25th min of exercise at a constant pace.
This principle has been held to be true for horses too
(Persson, 1983; Straub et al., 1984; Wilson et al., 1983;
Snow, 1987), even though the relationship has not been
demonstrated with the same degree of experimental
vigor. Thus, questions exist as to the application of this
principle in the horse. Therefore, the objective of this
study was to determine and validate maxLASS of Heck
et al. (1985) in horses resembling the experimental design as much as possible.
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Maximal lactate steady state in blood of horses
Table 1. Description of horses exercised continuously at their v2 and v4 (velocities run under the defined conditions
inducing a blood lactate concentration of 2 and 4 mmol/L, respectively) for 15 to 30 min
Horse
Track
Use
A
B
Trot races
Trot races
Trot races
Gallop races
Gallop races
Pleasure riding
Eventing LL1
Show jumping and dressage LL
Eventing LL
Eventing LL
Pleasure riding
Trot races
Trot races
Endurance 160 km
C
D
E
Age, yr
v4, m/s
v2, m/s
Speed run,
m/s
Duration of
exercise, min
7
3
4
7
7
6
8
8
8
6
6
4
4
11
10.03
10.33
9.67
9.30
9.63
6.83
9.17
8.00
7.83
9.17
7.75
10.25
9.25
7.17
9.12
9.67
7.40
8.00
8.00
6.00
7.28
6.75
6.58
7.17
6.62
8.33
7.33
6.50
9.12
9.67
7.40
8.00
8.00
6.00
7.17
6.67
6.67
7.17
6.33
8.15
7.33
6.00
30
20
30
15
20
20
20
25
25
20
25
25
25
20
1
LL = light level.
MATERIALS AND METHODS
This experiment was approved by the Animal Welfare Committee of the State of Baden-Württemberg,
Germany.
Horses and Study Design
For the maxLASS determination and validation in
the laboratory, 4 Thoroughbred horses were used (mean
BW = 451 ± 21 kg). One was 3 yr old, and 3 were 4
yr old (1 gelding, 3 mares). The horses were housed
in 3- × 3-m boxes in a barn. The daily feeding ration
consisted of 4.5 kg of concentrate, 0.2 kg of mineral
supplement, and 5 kg of silage (grass:corn, 2:1). On
days with exercise, the horses received an additional 1
kg of concentrate. Hay, straw, and water were available
ad libitum. The horses used were very well adapted to
run on a treadmill and all other handling procedures,
having participated previously in a conditioning study
for 11 mo (Werkmann et al., 1996). During this period,
horses were exercised almost constantly at intensities of
v2.5 or v4 for 5, 15, or 25 min with 1 d of rest between
exercise days. The exercise tests and workouts were
done on a high-speed treadmill with an incline of 3%
(Mustang, Kagra AG, Fahrwangen, Switzerland).
The results of the maxLASS determination and validation study in the laboratory were corroborated under
field conditions with 14 horses used in different sport
disciplines (Table 1).
Standardized Exercise Tests
MaxLASS Determination in the Laboratory.
The speeds to exercise the horses continuously for 40
min at v1.5, v2, v2.5, and v3 were calculated for each
individual, using an exponential equation that fit the
BLRS relationship obtained with a SET performed by
the horses at the start of the study.
The following formula was applied to calculate the
speeds to exercise the horses (Galloux, 1991): [LA] =
e(Av+B) + C, where [LA] = blood lactate concentration
in mmol/L, v = speed in m/min, e = 2.1828, A = coefficient of curvilinearity, B = constant, C = constant.
After calculation of the coefficients A, B, and C (using
SPSS/PC 4.01, SPSS Inc., Chicago, IL) v1.5, v2, v2.5,
and v3 could be determined using the equation v =
(ln([LA] + C) − B)/A.
The protocol for the SET started with a warm-up at
0% incline for 5 min at 1.6 m/s, followed by a 5-min
walk at 3.4 m/s. Thereafter, the slope of the treadmill
was set at 3% for the SET with several steps of 5-min
duration. Between 2 consecutive steps, horses walked
60 s. The SET was finished when the blood [LA] of the
horse reached 4 or more mmol/L. The speed in the first
step was 6.0 m/s, and it was increased in each consecutive step by 0.5 m/s. The increase in speed in each step
was such that a continuous increase of the blood [LA]
from the concentration before exercise but after warmup to 4 or more mmol/L was obtained in not less than
4 steps.
MaxLASS Determination in the Field. The
SET prescriptions varied according to the length and
surface of the field track as well as to the use of the
horses involved (Table 2). The number of laps run in
each interval was set to have a total duration of at least
5 min/interval (the term interval is used because the
duration of exercise was defined by a distance to be run
as opposed to a step where the duration of exercise is
defined by a time to be run). After the fourth interval,
the duration could be shorter than 5 min when an additional lap would have meant much more work for a
horse (Table 2). The increase in speed for each interval
was such that in all conditions, a continuous increase
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Lindner
Table 2. Track description and standardized exercise test (SET) prescriptions
Track
SET
Name
Surface
Length, m
A
Gravel
1,264
B
Gravel
1,200
C
Sand
1,912
D
Sand
600
E
Gravel
900
Interval
1st
2nd
3rd
4th
1st
2nd
3rd
4th
5th
1st
2nd
3rd
4th
5th
6th
1st
2nd
3rd
4th
5th
6th
1st
2nd
3rd
4th
5th
No. of
laps run
2
3
3
2
2
2
2
3
3
1
1
1
1
1
1
2
3
3
4
4
4
2
2
3
3
3
(2)1
(3)1
(3)1
(4)1
(4)1
Prescribed
speed, m/s
7.3
8.7
10.0
11.4
6.0
7.5
8.8
10.0
10.8
6.0
7.0
8.0
9.0
10.0
11.0
4.0
5.0
6.0
7.0
8.0
9.0
6.5
7.2
8.3
9.2
10.0
(5.0)1
(5.8)1
(6.7)1
(7.5)1
(8.3)1
1
Numbers in parentheses are for endurance horses, and outside numbers are for Standardbreds.
of the blood [LA] from the concentration measured before exercise but after warm-up to 4 or more mmol/L
was obtained in not less than 4 steps. The speeds were
known because the horses were tested regularly to guide
their training (Trilk et al., 2002). It was achieved in all
horses but one: the 6-yr-old pleasure riding Warmblood
tested on track D already had a blood [LA] of more
than 4 mmol/L after 3 intervals (Table 1). After each
interval, horses were stopped for 1 min in most cases,
but in some cases for 3 min, because of the regulations
prevailing at the track (i.e., had to take horse out of
track: tracks A, B, and E). The v2 and v4 were calculated using the formula mentioned previously (Galloux,
1991).
Exercise
MaxLASS Validation in the Laboratory.
Within 2 wk after the SET, each horse performed several exercises over 40 min at constant paces. Horses
always had at least 72 h of recovery between exercise
sessions. Horses were first submitted to the exercise at
v2.5. Thereafter, because of the results, the horses were
not submitted to exercise at their v3 but v2 and then
v1.5. During the exercise, horses were fitted with a commercial heart rate meter to record their heart rate at
5-s intervals (Polar Sport Tester, Kempele, Finland).
MaxLASS Validation in the Field. The day after SET, horses ran continuously at their v2 or a speed
slightly below or above between 15 and 30 min depending on the decision of the rider or trainer (Table 1).
Blood Sampling and Handling
MaxLASS Determination in the Laboratory.
Blood (5 mL) was sampled by venipuncture from a jugular vein as soon as possible after the end of each step,
normally within 10 s. The blood was collected into Naheparinized evacuated tubes (Vacutainer, Becton Dickinson, Heidelberg, Germany). Then 20 µL of blood was
collected into a disposable capillary pipette (Blaubrand
intramark, Brand Cat. No. 7087181, Wertheim, Germany) and applied on a test strip of a pocket analyzer to
obtain within 1 min the blood [LA] and stop the SET
if the value was above 4 mmol/L. Another 20 µL of
blood were immediately transferred to vials containing
200 µL of ice-cold 0.6 N perchloric acid. The supernatant was centrifuged at 20 to 25°C for 5 min at 12,000
× g (Biofuge A Heraeus-Christ, Osterode, Germany),
transferred to another vial, and kept frozen at −20°C
until analysis.
MaxLASS Determination in the Field. Blood
samples were always obtained within 20 s of finishing an
interval by puncturing a jugular vein and collecting the
samples in Na-heparinized evacuated tubes. The blood
[LA] was measured immediately with a pocket analyzer
(Accusport, Boehringer Mannheim, Germany).
2041
Maximal lactate steady state in blood of horses
MaxLASS Validation in the Laboratory. Venous blood was collected before the beginning of each
exercise session after warm-up (5 min at 1.6 m/s and 5
min at 3.4 m/s), every min during the first 6 min of exercise and then every 2 min thereafter until the end of
the exercise. Blood was sampled into a syringe from an
indwelling catheter (Braunüle MT 14G, No. 420 638/0,
B. Braun, Melsungen, Germany) secured in the right
or left jugular vein alternatively from SET to SET; a
140-cm section of tubing (Heidelberger Verlängerung,
No. 0409 7408, B. Braun) was attached to the catheter.
Patency of the catheter and tubing was maintained by
flushing 10 mL of 0.9% saline into the catheter after
obtaining the blood sample. Saline remaining within
the catheter was removed by withdrawing and discarding 20 mL from the catheter before collecting the
sample. The venous sample in the syringe was poured
into heparinized vials (Vacutainer, Becton Dickinson,
Heidelberg, Germany). The samples were handled as
described before for the samples obtained for the maxLASS determination in the laboratory.
MaxLASS Validation in the Field. Blood
samples were obtained by puncturing a jugular vein (5
mL) and collected in Na-heparinized evacuated tubes
at 5-min intervals between the 10th and 30th min of
exercise. For [LA] analysis, the portable lactate analyzer was used.
Blood Analysis
Two systems for lactate analysis were used: 1) a
pocket analyzer (Accusport, Boehringer Mannheim,
Germany), and 2) an EPOS 5060 lactate analyzer (Eppendorf, Wesseling, Germany) according to the method
of Noll (Bergmeyer, 1974), using an enzymatic test kit
(Boehringer Mannheim No. 1178 750, Mannheim, Germany).
The values obtained with Accusport were used to decide when to terminate the SET of the horses in which
maxLASS was determined (laboratory and field) and
to measure the blood samples obtained during the field
maxLASS validation study. The values measured with
the EPOS 5060 lactate analyzer were used to determine
and to validate maxLASS in the laboratory.
Determination of maxLASS as Postulated
by Heck et al. (1985)
The same definition of Heck et al. (1985) was adopted: maxLASS is the maximal speed at which the blood
[LA] does not change by more than 1 mmol/L between
the 5th and 26th min (1 min more than the original
definition) of continuous exercise.
Data Analysis
Mean and SD (x ± SD) are used to describe the data
(Statview, SAS Inc., Cary, NC). Data were analyzed
using a t-test for paired comparisons with P < 0.05 accepted as significant.
RESULTS
MaxLASS Determination and Validation
in Laboratory
The mean v1.5, v2, v2.5, and v3 of the horses was 8.1
(±0.5), 8.9 (±0.5), 9.3 (±0.5), and 9.6 (±0.5) m/s,
respectively. In the horses exercising at their v1.5, the
blood [LA] differed by more than 1 mmol/L from the
value in the 5th min after the 26th min or later, exercising at their v2 between the 14th and 18th min, and
exercising at their v2.5 between the 12th and 20th min
of exercise (Figure 1). After 40 min of exercise, the
mean blood [LA] of the horses running at their v1.5
was 4.9 ± 0.14 mmol/L, running at their v2 it was 5.5
± 1.47 mmol/L, and running at their v2.5 it was 8.5 ±
2.47 mmol/L (P < 0.05 between all comparisons). The
heart rate of the horses exercising at v1.5 were 155 ± 7
beats/min, at v2 167 ± 10 beats/min, and at v2.5 170
± 5 beats/min.
MaxLASS Determination and Validation
in the Field
There was no difference between the v2 of the horses
and the speed run during exercise (P > 0.05; v2 of 7.48
± 1.04 m/s, speed run 7.22 ± 0.96 m/s). The mean
blood [LA] did not vary by more than 1 mmol/L in the
horses during the time that they were exercised continuously at their individual v2 (Table 3).
DISCUSSION
The results of this study showed that for horses, the
concept of v4 as the point of the BLRS curve indicating
the aerobic-anaerobic lactate threshold, and therefore
maxLASS, is not valid. Running the horses at their
v2.5 and v2 elicited an increase of the blood [LA] by
more than 1 mmol/L before 25 min of continuous exercise. The different behavior of the blood [LA] for the
3 speeds run was also clearly shown after 40 min of
exercise, when the greatest values were measured for
the exercise at v2.5, the least for exercise at v1.5, and the
intermediate mean blood [LA] at v2. These observations
demonstrate that the v1.5 was the intensity which best
fulfilled the definition of maxLASS reported by Heck
et al. (1985).
The experiment performed in the field to validate
the findings under laboratory conditions on a treadmill
demonstrated that there is a large likelihood for horses to maintain a constant blood [LA] during at least
20 min of continuous exercise when they run at their
individual v2, provided v2 is determined with a SET
prescription like the one used in this study. The blood
[LA] values of these horses were measured with a por-
2042
Lindner
table lactate analyzer. This portable analyzer supplies,
within the range of values measured, values greater by
about 0.5 mmol/L than the automated analyzer used
to measure the samples obtained of the horses on the
treadmill (Lindner, 1996). Thus, the values measured
between both methods are not the same but can be
compared.
The results of the actual study have to be compared
with those of earlier studies (Valette et al., 1989, 1993;
Bourgela et al., 1991; Gottlieb-Vedi et al., 1994; Persson
et al., 1995). None of those authors reported the maxLASS at v4. These results are not surprising because
the SET applied in all of those studies did not conform
to the SET prescription of Heck et al. (1985). Heck et
al. (1985) examined their athletes with an incremental
SET that had steps of 5-min duration and had to have
at least 4 steps before the athletes had a blood [LA]
of 4 mmol/L or more. The SET used by Valette et al.
(1989, 1993) and Bourgela et al. (1991) had steps with
a duration of 3 min, whereas the SET of Gottlieb-Vedi
et al. (1994) and Persson et al. (1995) had steps of
2-min duration only. At the same speeds, longer steps
will induce greater blood [LA], and these will reduce
the variables derived mathematically from the BLRS
curve, like v4 (Köster, 1996). Thus, a velocity producing a given [LA] calculated in the present study was less
than that determined in the studies mentioned.
Table 3. Development of the blood lactate concentration of horses during continuous exercise in the field
at their v2 (velocity run under the defined conditions
inducing a blood lactate concentration of 2 mmol/L; 14
horses; mean ± SD)
Time of blood sampling
during exercise
After
After
After
After
After
10
15
20
25
30
min
min
min
min
min
Blood lactate
concentration,
mmol/L
1.63
1.75
1.84
1.54
1.60
±
±
±
±
±
0.36
0.58
0.55
0.25
0.71
No. of
horses
14
13
13
7
2
In 5 horses run between 25 and 33 min, according to
their lactate minimum speed, Gondim et al. (2007) found
that the blood [LA] stayed constant during the continuous exercise at concentrations less than 2 mmol/L. The
prescription used by Gondim et al. (2007) to determine
maxLASS is very different than the prescriptions used
by Heck et al. (1985) and in this study, but it is remarkable that when the horses ran at a speed 10% above
their lactate minimum speed, the blood [LA] stayed
at the initial concentration only for about one-fifth of
the time than when they were exercised at their lactate
minimum speed.
Figure 1. Development of the blood lactate concentration of horses during continuous treadmill exercise at speeds derived from a standardized exercise test (4 horses; mean ± SD; v1.5, v2, v2.5 = velocities run under defined conditions inducing 1.5, 2, and 2.5 mmol/L blood lactate
concentrations).
Maximal lactate steady state in blood of horses
The maxLASS of running horses determined in this
study was less than the maxLASS of running man
(Mader et al., 1976; Heck et al., 1985). This may be a
consequence of the larger proportion of the total skeletal muscle mass involved in the exercise. This conclusion can be drawn reviewing the maxLASS studies done
with athletes doing different types of physical activities.
In cycling, the maxLASS of the athletes were, on average, at a blood [LA] of 4.3 mmol/L (Heck and Rosskopf,
1994), in rowing athletes at 3 mmol/L (Beneke, 1995),
in canoeing athletes at 6.2 mmol/L (Krüger et al.,
1990), and in skating athletes at 6.6 mmol/L (Beneke
et al., 1991). These results show that the larger the
proportion of the total skeletal muscle mass used for
exercise, the less the maxLASS. This was demonstrated
experimentally by Beneke et al. (2001). Differences in
blood distribution during exercise, muscle respiratory
capacity, and fiber types engaged during different types
of motor activity may be other factors affecting the
maxLASS (Ivy et al., 1980).
The v4 has long been used to compare sport horses
and examine the effect of training (Thornton et al.,
1983; Wilson et al., 1983; Straub et al., 1984; Auvinet
et al., 1991; von Wittke et al., 1994). Also, good relationships have frequently been described between v4
and variables of performance such as earnings, placings,
and winnings (Demonceau, 1989; Erickson et al., 1990,
1991; Galloux, 1991; Casini and Greppi, 1996; Couroucé, 1997; Ponchard, 1998; Lindner, 2010). Only more
recently have studies been done to examine the effects
of conditioning horses with the speed of exercise guided
by different blood [LA] obtained with a SET of similar
prescription as in this study (Werkmann et al., 1996;
Gansen et al., 1999; Trilk et al., 2002; Lindner et al.,
2009). The largest effect on the increase of v4 so far has
been observed exercising horses at their v2 for 45 min
(Trilk et al., 2002). Future studies will show whether
guiding the exercise speed with other variables of the
BLRS in combination with different duration and frequency of exercise are more effective.
In conclusion with exercise at v1.5 the blood [LA] of
horses stayed within 1 mmol/L of difference during 20
min of continuous exercise after 5 min running at the
same pace. Hence, for horses, not v4, but much less
blood LA-derived speeds best predict maxLASS as defined by Heck et al. (1985) and therefore the aerobicanaerobic threshold.
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