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International]ournal of Sport Nutrition, 1 996,6, 295-306

O 1996 Human Kinetics Publishers, Inc.

Branched-Chain Amino Acid Supplementation


During Repeated Prolonged Skiing Exercises
at Altitude

Andr6 X. Bigard, Pierre Lavier, Lionel Ullmann,


H6lGne L egrand, Philippe Douce, and Charles Y. Guezennec
This study was conducted to test the hypothesis that a branched-chain amino
acid (BCAA) supplementation would minimize changes in body composition
and alterations in plasma amino acid profile induced by prolonged exercises at
altitude. Twenty-four highly trained subjects participated in six successive
sessions of ski mountaineering (6-8 hr duration, altitude 2,500-4,100 m).
Twelve subjects took a dietary supplement of BCAA (BCAA group) and 12
took a dietary supplement that was 98% carbohydrate (C group). Body weight
decreased in C subjects (-2.1%, p < .01), while the body weight loss recorded
in the BCAA group was not statistically significant (-1.2%, NS). Changes in
body composition that resulted from repeated skiing exercise at altitude were
not significantly minimized by BCAA administration. Peak power output
recorded during an incremental bicycle exercise decreased in C subjects but did
not change significantly in BCAA subjects. Results of this study demonstrate
that neither changes in body composition related to the ski mountaineering
program nor muscular performance during isometric contraction was signifi-
cantly affected by BCAA administration.

Key Words:repeated endurance exercise, body composition, hormones, plasma


amino acid profile

Energy balance studies in athletes engaged in low- to moderate-intensity


exercises of long duration (50 to 70% of the maximal oxygen uptake, or VO,max,
2 to 6 hr - day-') have indicated that ultraendurance training may require an increase
in protein intake above the Recommended Dietary Allowances (RDA) (for review,
see 19). Under conditions of reduced substrate availability, depressed protein
synthesis and/or elevated protein degradation may provide a significant quantity of
amino acids for use as an auxiliary fuel (7). The rate of amino acid oxidation is
increased during prolonged exercise (23), but only a few amino acids are oxidized
at a significant rate, among which are branched-chain amino acids (BCAA; i.e.,

A.X. Bigard, L. Ullmann, H. Legrand, P. Douce, and C.Y. Guezennec are with the
D$artment de Physiologie Systernique, CERMA, B.P. 73, 91223 Bretigny-sur-Orge,
France. P. Lavier is with the Ecole Militaire de Haute Montagne, 74400 Charnonix, France.
296 1 Bigard, Lavier, Ullmann, et al.

leucine, isoleucine, valine), alanine, glutamate, and aspartate (10). Some evidence
exists that oxidation of BCAAs is most obvious in well-trained subjects (14).On the
other hand, urea production has been reported to be greater when glycogen stores
are low at the onset of exercise (21), and it has been hypothesized that an adequate
caloric intake prevents N-excretion during prolonged exercise (30).
Evidence exists that plasma BCAA levels decline during endurance or
ultraendurance exercise (2,3,20,25). It has been hypothesized that these changes
in the plasma amino acid profile could increase concentrations of 5 hy-
droxytryptamine (5-HT) in the brain (22), which is thought to partly contribute to
fatigue during prolonged exercise. However, there is little information concerning
the cumulative effects of repeated daily sessions of prolonged exercise on amino
acid oxidation and plasma amino acid concentration. We previously reported
that repeated prolonged exercise at altitude reduced plasma BCAA concentra-
tions measured at rest by 14 to 22% (2) and that a high protein intake (2.5 g . kg-'
day-') minimized the decrease in plasma BCAA levels but reduced the isometric
endurance capacity of the knee extensor muscles. This alteration in muscle
performance was probably related to reduced carbohydrate intake and decreased
muscle glycogen stores.
High altitude exposure is known to decreasebody weight in those accustomed
to lower altitudes (4,9, 12). Weight changes are paralleled with reductions in fat
mass and/or muscle mass. The magnitude of these anthropometricchanges depends
on the altitude and duration of exposure. Because loss of muscle mass may
negatively influence exercise capacity, an attempt has been made to minimize the
altitude-inducedchanges in body composition.In this respect, it was recently shown
that a dietary supplementationof BCAA minimized muscle loss during trekking at
high altitude (24).
In the present study, it was hypothesized that daily administration of BCAA
in the diet during repeated prolonged exercise at high altitude would prevent an
altitude-related alteration in body composition and limit alterations in plasma
amino acid profile without affecting skeletal muscle performance. To test these
hypotheses, we designed a study in which one group of highly trained subjects
participated in six successive sessions of ski mountaineering while receiving a
dietary supplement of BCAA.

Methods
Subjects
Twenty-four healthy male volunteers (25-42 years old) gave their informed written
consent to participate in this study. The experimentalprotocol was approved by the
Cochin-Port Royal HospitalEthics Committee.All subjects had been ski mountain-
eering for many years and were highly trained in this type of exercise. This
experiment was conducted in the field, and exercise conditioning was very similar
to that previously described (2). The subjects participated in six successive sessions
of ski mountaineeringin the French Alps. Before departure, subjects were randomly
assigned to one of two treatment groups: One group received a dietary supplement
of carbohydrates (C, n = 12) and the other a supplement of branched-chain amino
acids (BCAA, n = 12).
Anthropometric measurements, determination of physical performance, and
blood sampling were performed in the laboratory before and immediately after the ski
Branched-ChainAmino Acid 1297

mountaineeringprogram. Each stage of the ski mountaineeringprogram consisted of


7-9 hr of walking on skis with a backpack (15-20 kg), at altitudes ranging between
2,500 and 4,100m. The skimountaineeringexercisesstartedbetween7 and 8 a.m., after
a hearty breakfast. The subjects walked on skis until they reached the refuge. They
stopped every hour for a 5-min break to drink water and ingest foodbars. As soon as
they reached the refuge, the subjects had a substantial meal, followedby a dinner at 7-
8 p.m. Daily ascents ranged between 1,200 and 2,100 m.

Dietary Protocol
During the ski mountaineeringperiod, the subjects consumed a standard prepackaged
diet to which different dietary supplements were added according to the experimental
protocol. Diets were distributed to the subjects on the first day of the skiing period and
were consumed up to the time of the last tests after subjectsreturned to the laboratory.
Diets were isocaloric (4,000 kcal . day', 16,760kJ - day'), the standard prepackaged
diet providing 3,200 kcal - day' (13,410 kJ - day') and the supplement 800
kcal . day1(3,350 kJ . day1).Although no measurements could be made during ski
mountaineering, it is likely that energy expenditure slightly exceeded energy intake.
However,it has recently been shown that a moderate deficitbetween energy intakeand
expenditure did not affect performance after heavy military maneuvers and that diets
providing only 3,200 kcale day' had no deleterious effect on physical performance
compared to diets providing 4,200 kcal - day1(11).
In addition to the prepackaged diet, subjects of the C group consumed a
dietary supplement composed of 98% carbohydrates. The final macronutrient
composition of the diet provided -58% of calories as carbohydrates, -31% as fat,
-
and 11% as protein (1.2 g .kg-' - day'). Each subject in the BCAA group consumed
a dietary supplement containing a mixture of BCAA (7.8 g . day-' leucine,
3.4 g . day*isoleucine, and 11.2 g . day-' valine, i.e., 35% leucine, 15% isoleucine,
and 50% valine). The composition of the dietary supplement for the BCAA group
was selected accordingto the beneficial effects of such a mixture on both mental and
physical performance during prolonged exercise (3). This dietary supplement
provided -49% of energy from carbohydrates, -38% from fat, and -13% from
protein. Subjects in the BCAA group were given a total diet that provided -54% of
energy from carbohydrates, -32% from fat, and -14% from protein (1.44 g . kg-'
day-'). In both groups, diets exceeded recommended dietary allowancesfor protein
(0.8 g - kg-' . day-'). The subjects were advised to consume the dietary supplement
while walking on skis. Although dietary compliance was not assessed in the present
study, the experimental conditions contributed to maximize subjects' cooperation:
Two investigatorsparticipated in the ski mountaineering program, all subjects were
active mountaineers who were highly motivated by this experiment, and isolation
in the mountains during the 6 days of the ski trek ensured that subjects did not
consume food from any source other than the distributed diets. Consumption of the
dietary supplements was verified each evening.

Anthropometric Measurements
Before and immediately after the ski mountaineering period, body weight (BW) was
measured on the same standard medical scale with an accuracy of +lo0 g, while the
subjects wore the same shorts. The anthropometric assessment consisted of four
skinfold thickness and one circumference measurement obtained while subjects
298 / Bigard, Lavier, Ullmann, et a/.

were standing. One trained investigator made all anthropometric measurements.


Percentage of body fat (%BF) was calculated from triceps, biceps, subscapular, and
suprailliac skinfolds using a Harpenden calliper and using the tables from Durnin
and Womersley according to the subjects' ages (8). The anthropometric measure-
ments were obtained on the left side of the body. Triceps skinfold thickness and
circumferences of the middle of the upper arm were used to estimate bone free arm
muscle area (AMA) according to the formula provided by Heymsfield et al. (16):
{[muscle arm circumference - (IT * triceps skinfold)12/ 47r) - 10

Exercise Testing
Subjects' maximal aerobic capacity (maximal oxygen uptake, V0,max) and muscle
performance were measured 2 days before the ski mountaineering program and
within 1 day after returning to the laboratory. At the time of the study, all subjects
were in a maintenance phase of their skiing training program. Two days elapsed
between the last bout of ski mountaineeringexercise and the pretrek measurements.
The V0,max was determined during an incremental protocol on a mechanically
braked cycle ergometer (Ergomeca, SOREM, Toulon, France). The maximal exercise
test consisted of a 3-min warm-up at 120W, followed by aprogressiveresistive phase
during which the workload was increased by 22.5 W . min-'. A pedaling frequency of
75 rpm was selected. Oxygen uptake was measured using an automated gas analysis
system (SensorMedics 2900, SensorMedicsCorporation, Yorba Linda, CA).
Isometric knee extension force was measured on the right leg, with subjects
comfortablyseatedin an adjustablechair. Measurements were made on an isokinetic
dynamometer (Cybex I1 equipment, Cybex Division of Lumex, Inc., Ronkonkoma,
NY). The maximum isometric contraction (MVC) that could be produced by the
knee extensor muscle group was determinedwith the hip flexed to 100" and the knee
in 80" flexion. MVC was defined as the maximal force the subject could sustain for
1 s and was determined from at least four contractions. Approximately 2-3 min
elapsed between two maximal contractions. Fifteen minutes after determination of
MVC, subjects were required to sustain to the point of exhaustion a contraction at
constant force correspondingto 50% MVC. Endurance time was defined as the time
from the start of contraction until exhaustion, and exhaustion was defined as the
point at which the subject was unable to maintain the required tension. All exercise
capacity measurements were performed between 3 and 7 p.m.

Blood Sampling and Analyses


Blood was sampled from a forearm vein using an indwelling catheter. A sample of
20 ml blood was obtained at rest in the overnight fasted state (between 6 and 6:30
a.m.), before the ski mountaineering sessions, and within 12 hr after completion of
the ski mountaineering sessions.
The hematocrit was determined immediately by high-speed centrifuge. Each
blood sample (10 ml) was deproteinized with 8% perchloric acid, or plasma was
separated from the remaining blood by centrifugation and divided into separate
aliquots. Samples were quickly frozen with liquid nitrogen and stored at -20 "C for
further analyses. Assays were performed within 30 days. Lactate, glucose, 3-
hydroxybutyrate, and glycerol were assayed in neutralized samples using enzy-
matic methods (1). Serum free fatty acids (FFA) were determined according to the
Branched-ChainAmino Acid / 299

method of Ho (17). The coefficients of variation (CV) of these assays were less than
5% for concentrations in the physiological range. Plasma ammoniumconcentration
was determined using a colorimetric method with an Ektachem DT60 analyzer
(Eastman Kodak Company, Rochester, NY). The CV was 2.8% and 5.4% for
two levels of standard (200 pmol - L-' and 588 ymol L-', respectively). Hor-
mone determinations were made in duplicate using commercially available radio-
immunoassay kits (CIS Biointernational, Gif-sur-Yvette, France): insulin
(SB-INSI-5) (CV 8 and 4% for two levels of standard, 18.2 and
36.5 mUI - L-I,respectively); cortisol (SB-CORT) (CV 5 and 4% for two levels of
standard, 35 and 125 ng - ml-', respectively); testosterone (SB-TESTO) (CV 8 and
5% for two levels of standard, 2.4 and 9.33 ng . ml-', respectively); and growth
hormone (GH) (SB-HGH) (CV 9 and 7% for two levels of standard, 1.9 and 5.2 ng
- ml-', respectively).
Plasma amino acid concentrations were determined at rest, under overnight-
fasted conditions, by reversed-phase, high-performance liquid chromatography
according to the method described by Henriksson and Meredith (15). Prior to
separation of amino acids, plasma samples were deproteinized by ultrafiltration.
Derivatization was performed with phenylisothiocyanate(PITC, 20 p1) in solution
with methanol (140 yl), triethylamine (20 pl), and water (20 PI). For greater
accuracy, L-nomaline was used as an internal standard.

Statistics
+
All values are presented as means standard error of the mean (SEM). Both main
effects-treatment (BCAA vs. C) and time (pre- vs. post repeated exercises)-and
the interaction of treatment and time were analyzed with a two-way analysis of
variance. Where appropriate, a Student's t test for paired samples was used to
determine significant differences between pre- and post repeated skiing exercises
within each group (C and BCAA). P values of <.05 were considered significantafter
adjustment using the Bonferroni correction.

Results
Anthropometric data are shown in Table 1.The subjects of the C group lost an average
of 1.55 -t 0.48 kg (-2.1%, p < .01) during the 6-day trek. There was a nonsignificant
mean loss of 0.8 kg in the BCAA group. After the ski mountaineering exercise, %BF
decreased by 11.8%( p < .01), accounting for 77% of total weight loss in the C group.
In theBCAAgroup, there was a 13.7% decreasein %BFfrompre- to posttrek ( p < .Ol),
accounting for 76% of the total weight loss. AMA estimated from skinfold and
circumference measurements was not significantly altered by the skiing exercise
program.
Exercise capacity data are given in Table 2. Maximal oxygen uptakeremained
unaffected by the repeated ski mountaineering exercises. The power output mea-
sured at V0,max (Wmax) was significantlyaltered by the repeated skiing exercises.
At the end of the ski trek, Wmax had decreased in C subjects (-9%,p < .05) and had
not significantlychanged in the BCAA group (-2.5%, NS). Neither the peak torque
measured during the maximal voluntary isometric contraction of the knee extensor
muscle group (MVC) nor the endurance time at 50% MVC was significantly
affected by the repeated skiing exercises (Table 2).
300 / Bigard, Lavier, Ullmann, et al.

Table 1 Anthropometric Measurements Before and After the Successive


Ski Mountaineering Stages

C group (n = 12) BCAA group (n = 12)


M SEM M SEM
- - - -

B w (kg) Pre 71.72 2.63 70.18 1.47


Post 70.17* 2.42 69.36 1.47
%BF (%) Pre 10.33 1.02 8.09 0.61
Post 9.11* 1.02 6.98* 0.53
AMA (cm2) Pre 48.07 1.81 46.61 1.99
Post 48.33 1.64 46.57 1.53

Note. Pre = before the repeated ski mountaineering exercises; Post = after the repeated
exercises; BW =body weight; % BF = percentage body fat; AMA = bone-free arm
muscle area. Significant differences compared with values measured before the ski
mountaineering exercises, *p < .01.

Table 2 Exercise Capacity Data

C group (n = 12) BCAA group (n = 12)


M SEM M SEM

90,max (ml . min-' . kg-') Pre


Post
Wmax (W) Pre
Post
MVC (N . m) Pre
Post
Endurance time (s) Pre
Post

Note. V0,max = maximal oxygen uptake; Wmax = maximal workload during incremen-
tal bicycle exercise; MVC = peak torque during maximal isometric voluntary contraction
of the quadriceps femoris muscle. Significant differences compared with values measured
before the ski mountaineering exercises, *p < .05.

Table 3 provides the plasma metabolic profile. Plasma volume was es-
timated by the packed cell volume. No significant changes occurred in hema-
tocrit after the repeated skiing exercises. Plasma insulin concentrations at rest
were significantly decreased after the ski mountaineering program (p< .05), especially
in the BCAA group (-26%, p < .05).There was an increase in blood concentrations of
P A (p < .01) that was significant in both C (p < .01) and BCAA @ < .05) groups.
Hormonal changes are shown in Table 4. A significant decrease in plasma
cortisol concentration was recorded after the repeated skiing exercise in the C group
(p < .05). Plasma GH concentrations measured at rest were slightly increased after
the repeated sessions of ski mountaineering in the BCAA group (p < .03).
Branched-Chain Amino Acid / 301

Table 3 Plasma Metabolic Profile

C group (n = 11) BCAA group (n = 11)


M SEM M SEM

Hematocrit (%) Pre


Post
Glucose (mmol . L-I) Pre
Post
Insulin (mu - L-I) Pre
Post
Lactate (mmol .L-I) Pre
Post
FFA (mmol . L-I) Pre
Post
Glycerol (mmol . L-l) Pre
Post
P-hydroxybutyrate Pre
(pmol . L-I) Post
Ammonia (kmol . L-I) Pre
Post
- -

Note. FFA = free fatty acids. Significant difference with values measured before the ski
mountaineering exercise, *p < .05, **p < .01. For further details, see Table 1.

Table 4 Blood Hormone ConcentrationsMeasured at Rest Before and After the


Ski Mountaineering Program

C group (n = 11) BCAA group ( n = 11)


M SEM M SEM

Cortisol (ng . ml-I) Pre 118.78 21.52 79.36 16.93


Post 71.78* 10.05 91.05 17.61
Testosterone (ng . ml-I) Pre 6.02 0.52 6.23 0.39
Post 6.22 0.5 1 5.59 0.19
GH (ng . ml-') Pre 0.19 0.02 0.19 0.02
Post 0.29 0.06 0.78* 0.22

Note. GH = growth hormone. *p < .05. For further details, see Table 1.

Regarding plasma amino acid concentrations,analysis of variance of plasma


BCAA data showed a significantinteraction of treatmentby time (p < .01). Repeated
prolonged skiing exercise caused a statistically significant decrease in the plasma
level of the three BCAAs in the C group (-16%, p < .01) (Table 5). Leucine
decreased by 14% ( p < .05), isoleucine by 10% (NS), and valine by 19%(p < .05)
(Figure 1). BCAA supplementation entirely suppressed the decrease in plasma
leucine concentrations that occurred after the repeated skiing exercises in the C
302 / Bigard, Lavier, Ullmann, e t a / .

Table 5 Plasma Amino Acid Concentrations Measured at Rest

C group (n = 11) BCAA group (n = 11)


M SEM M SEM

Plasma BCAA Pre


Post
Glutamine Pre
Post
Glutamate Pre
Post
Alanine Pre
Post
Tryptophan Pre
Post
- - - -

Note. Values are expressed as pmol . 100ml-' . *p < .05; **p < .01. For further details, see Table 1.

c leucine isoleucine valine

l2 1

BCCA
0II
before

leucine
after

isoleucine
before

valine
-
after

D nli after before

Figure 1-Plasma branched-chain amino acid concentrationsat rest, measured before


and after completion of the repeated skiing exercises in control group (C), and in
subjects who consumed the BCAA supplement (BCAA). Data are means f SEM;n = 11
in each group; *p ~p -05.
after
Branched-Chain Amino Acid / 303

group, whereas isoleucine concentrations remained unchanged (Figure 1). The


plasma concentration of valine was significantlyincreased by BCAA supplementa-
tion (+15%, p < -05).
Resting plasma glutarnine concentrations were increased after completion of
the ski mountaineeringprogram in theBCAA group (+26%,p < .05, vs. +20%, NS,
in C subjects) (Table 5). No significant changes were found in plasma concentra-
tions of glutamate or alanine in either the C or the BCAA group (Table 5). Plasma
concentration of total tryptophan increased in the BCAA group (+36%, p < -05)
while remaining statistically unchanged in the C group (+la%, NS).

Discussion
The present study was designedto determinethe effects of BCAA supplementation on
body weight, body composition, performance, and plasma amino acid concentsations
after the completion of six successive skiing exercises of long duration (7-9 hr) at
altitude. It was shown that subjects of the C group lost an average of 1.55 kg, whereas
the slightdecreaseinBW recorded in the BCAA group was not statisticallysigntfkant.
Boyer and Blume reported a mean loss of 1.9kg duringthe approach march (up to 5,400
m) of a Himalayan expedition (4). Guilland and Klepping observed body weight
decrements of 1.5 kg on Day 8 of exposure to 4,800 m in climbers during a mountain
expedition (12). These studies also demonstrated that exposure to extreme altitude
induces higher losses of BW. Exposure to slightly higher altitudes but with probably
lower energy expenditure induced a loss of BW varying between 1.1and 1.8kg, similar
to that recorded in the present study (24). It was shown that muscle loss during high-
altitude exposure can be reduced by daily administration of BCAA (24). The experi-
mental data obtained in the present study did not show any muscle loss as estimated by
variations of the arm muscle cross-sectional area (calculated from skinfold and
circumferencemeasurements).This apparent discrepancy may be explainedby the fact
that the subjects did not live at altitudes higher than 2,000-2,500 m and transiently
reached altitudes higher than 3,500 m. The observation that loss of BW was mainly
related to loss of %BF was consistent with previously reported data (4,9). Thus, the
results do not provide evidence that repeated exercise of long duration performed at
high altitude induces significant loss of fat-free mass.
The decrease in maximal power output that occurred in C subjects after
repetitive skiing exercises was not observed in subjects who had consumed the
BCAA supplement. This confirms findings by Schena et al. (24), who showed that
the decrease in lower limb muscle power estimated by repeated maximal jump
during altitude acclimatizationcould be minimized by BCAA supplementation.We
have previously reported that a daily protein intake equal to 2.5 g . kg-' decreased
the endurance time of the knee extensor muscles at 50% MVC under similar
experimental conditions (2).This finding was partly related to a low carbohydrate
content in the diet, and a decrease in muscle glycogen availability was expected. In
the present study, 58% of calories were provided as carbohydrate in the C group, in
comparison with 54% in the BCAA group. Although muscle glycogen concentra-
tions were not measured, it appears unlikely that this small difference in carbohy-
drate intakes between experimental groups altered muscle glycogen availability.
Whether the slight increase in plasma GH concentration observed at rest in
BCAA subjects is physiologically relevant remains to be determined. In compari-
son with the specific effects of endurance exercise (28), changes in plasma GH
304 / Bigard, Lavier, Ullmann, eta/.

recorded in the present study 12 hr after the last ski mountaineering exercise were
so small that their physiological significance may be questioned. Repeated daily
skiing exercises at high altitude also decreased plasma cortisol concentrations,
which was probably related to the repetition of prolonged high-intensity skiing
exercises rather than to hypoxia, since hypoxia did not affect plasma cortisol at rest
(13, 27). The reason why this decrease was only observed in the control group
remains unknown. Most of the changes seen in plasma FFA, glucose, and insulin
concentrationshave been reported in humans duringprolongedexposure to altitude.
Previous reports indicated-that resting plasma glucose concentrations were not
significantly affected by altitude (2, 32), while plasma insulin decreased during
long-term exposure to hypoxia (26). It has been shown that these changes are
associated with an increase in plasma FFA measured at rest (2,26, 31).
The results of the present study confirmed that six successive prolonged
exercises at high altitude altered plasma BCAA concentrations measured at rest.
These data are in accordance with those previously reported under similar experi-
mental conditions (2). It has been shown that marathon races (3) and exercises of
longer duration, such as 100-km races, 8 hr of cycling and running, or ultratriathlons
(6, 20, 25), were associated with a decrease in plasma BCAA concentration
measured 10 min to 2 hr postexercise. The alteration in average plasma BCAA
concentrationobserved in the present study was similar to that previously measured
under similar experimental conditions (-23%) (2). These changes were within the
range of decreases in plasma BCAA concentrationsinduced by a marathon (-19%)
(3), a 100-km race (-32%) (6), and an ultratriathlon lasting more than 23 hr (-18%)
(20). Previously reported results tend to support the hypothesis that both the
metabolic use of BCAA and the plasma levels of these amino acids are related to
training; that is, well-trained individuals tend to have decreased plasma levels of
BCAAs (14).The subjects who tookpart in the present study were all highly trained
in ski mountaineering and in long-duration endurance exercise. They displayed
high maximal aerobic power, reflecting their endurance-trained status (59.11 f 1.85
ml min-' . kg-' in the C group vs. 58.92 f 1.93 rnl .min-' . kg-' in the BCAA group).
These observations agree with the hypothesis that alterations in plasma BCAA
could be related to the training status of subjects.
One interesting finding of the present study was that the BCAA dietary
supplementation suppressed the exercise-induced decrease in plasma BCAA. The
preventive effect of BCAA supplementation on the decrease in plasma BCAA
concentrations was related to the concentration of the experimental dietary supple-
ment; 35%leucine entirely suppressed the expected decrease, 15%isoleucine had
no effect on the exercise-induced decrease, aid 50%valine significantly increased
the plasma concentration of this amino acid. It has been previously shown that
BCAA administration during prolonged exercise improves both mental and physi-
cal performance (3) and alters the perception of fatigue during high-volume training
(19). However, the hypothesis that BCAA administration delays fatigue during
prolonged exercise has been disputed (5, 29). Performance during prolonged
submaximal exercise was not measured in the present study; we only noted that the
overall perception of effort was similar in C and BCAA groups.
In conclusion, the changes in body composition that resulted from repeated
skiing exercise at altitude were not significantly minimized by BCAA administra-
tion. BCAA supplementationprevented the decrease in maximal workload related
to the repetitive skiing exercises but did not affect muscular performance during
isometric contraction. On the other hand, the results of the present study show that
Branched-Chain Amino Acid / 305

BCAA supplementation compensated for the decrease in plasma BCAA concentra-


tions that occurred following six successive ski mountaineering exercises.

References
I. Bergmeyer, H.U. Methods of Enzymatic Analysis, H.U. Bergemeyer (Ed.). New York,
London: Academic Press, 1974.
2. Bigard, A.X., P. Satabin, P. Lavier, F. Canon, D. Taillandier, and C.Y. Guezennec.
Effect of protein supplementation during prolonged exercise at moderate altitude on
performance and plasma amino acid pattern. Eur. J. Appl. Physiol. 66:5-10, 1993.
3. Blomstrand, E., P. Hassmkn, B. Ekblom, and E.A. Newsholme. Administration of
branched-chain amino acids during sustained exercise-Effects on performance and on
plasma concentration of some amino acids. Eur. J. Appl. Physiol. 63:83-88, 1991.
4. Boyer, S.J., and F.D. Blume. Weight loss and changes in body composition at high
altitude. J. Appl. Physiol. 57: 1580-1585, 1984.
5. Davis, J.M., S.P. Bailey, J.A. Woods, F.J. Galiano, M.T. Hamilton, and W.P. Bartoli.
Effects of carbohydrate feedings on plasma free tryptophan and branched-chain amino
acids during prolonged cycling. Eur. J. Appl. Physiol. 6 5 5 13-519, 1992.
6. Decombaz, J., P. Reinhardt, K. Anantharaman, G. von Glutz, and J.R. Poortmans.
Biochemical changes in a 100 km run: free amino acids, urea, and creatinine. Eur. J.
Appl. Physiol. 41:61-72, 1979.
7. Dohm, G.L., E.B. Tapscott, and G.J. Kasperek. Protein degradation during endurance
exercise and recovery. Med. Sci. Sports Exerc. 19:s 166-S171, 1987.
8. Durnin, J.V.G.A., and J. Womersley. Body fat assessed from total body density and its
estimation from skinfold thickness: Measurements on 481 men and women aged 16 to
72 years. Br. J. Nutr. 32:77-97, 1974.
9. Fulco, C.S., R.W. Hoyt, J. Baker-Fulco, J. Gonzalez, and A. Cymerman. Use of
bioelectrical impedance to assess body composition changes at high altitude. J. Appl.
Physiol. 72:2181-2187, 1992.
10. Goldberg, A.L., and R. Odessey. Oxidation of amino acids by diaphragms from fed and
fasted rats. Am. J. Physiol. 223:1384-1391, 1972.
11. Guezennec, C.Y., P. Satabin, H. Legrand, and A.X. Bigard. Physical performance and
metabolic changes induced by combined prolonged exercise and different energy
intakes in humans. Eur. J. Appl. Physiol. 68525-530, 1994.
12. Guilland, J.C., and J. Klepping. Nutritional alterations at high altitude in man. Eur. J.
Appl. Physiol. 54517-523, 1985.
13. H W n e n , K., A. Pakarinen, M. AlCn, and P.V. Komi. Serum hormones during
prolonged training of neuromuscular performance. Eur. J. Appl. Physiol. 53:287-293,
1985.
14. Henriksson, J. Effect of exercise on amino acid concentrations in skeletal muscle and
plasma. J. Exp. Biol. 160:149-165, 1991.
15. Henriksson, R.L., and S.C. Meredith. Amino acid analysis by reverse-phase high-
performance liquid chromatography: Precolumn derivatization with phenylisothiocya-
nate. Anal. Biochem. 136:65-74, 1984.
16. Heymsfield, S.B., C. McNamus, J. Smith,V. Stevens, andD.W. Nixon. Anthropometric
measurement of muscle mass: Revised equations for calculatingbone-free arm muscle
area. Am. J. Clin. Nutr. 36:680-690, 1982.
17. Ho, P.J. Radiochemical assay of long chain fatty acids using 63 N as tracer. Anal.
Biochem. 36:105-113. 1970.
306 / Bigard, Lavier, Ullrnann, eta!.

18. Kreider, R.B., and C.W. Jackson. Effects of amino acid supplementationon psychologi-
cal status during an intercollegiateswim season. Med. Sci. Sports Exerc. 26:s 115,1994.
19. Kreider, R.B., V. Miriel, and E. Bertun. Amino acid supplementation and exercise
performance. Analysis of the proposed ergogenic value. Sports Med. 16:190-209,1993.
20. Lehmann,M., M. Huonker, F. Dimeo, N. Heinz, U. Gastmann,N. Treis, J.M. Steinacker,
J. Keul, R. Kajewski,and D. Haussinger. Serum amino acid concentrationsin nine athletes
before and after the 1993 Colmar ultra triathlon. Int. J. Sports Med. 16:155-159, 1995.
21. Lemon, P.W.R., and J.P. Mullin. Effects of initial muscle glycogen levels on protein
catabolism during exercise. J. Appl. Physiol. 48:624-629, 1980.
22. Newsholme, E.A., I.N. Acworth, and E. Blomstrand. Amino acids, brain neurotransmit-
ters and a functional link between muscle and brain that is important in sustained
exercise. In Advances in Myochemistry, G. Benzi (Ed.). London: John Libbey Eurotext,
1987, pp. 127-133.
23. Rennie, M.J., R.H.T. Edwards, S. Krywawych, C.T.M. Davies, D. Halliday, J.C.
Waterlow, and D.J. Millward. Effect of exercise on protein turnover in man. Clin. Sci.
61:627-639, 1981.
24. Schena, F., F. Guenini, P. Tregnaghi, and B. Kayser. Branched-chain amino acid
supplementation during trekking at high altitude. Eur. J. Appl. Physiol. 65:394-398,1992.
25. Stein,T.P.,R.W. Syt,M. O'Toole,M.J. Leskiw,M.D. Schulter,R.R. Wolfe, and W.D.B.
Hiller. Protein and energy metabolismduring prolonged exercise in trained athletes. Int.
J. Sports Med. 10:311-316, 1989.
26. Stock, M.J., C. Chapman, J.L. Stirling, and I.T. Campbell. Effects of exercise, altitude,
and food on blood hormone and metabolite levels. J. Appl. Physiol. 45:350-354, 1978.
27. Sutton, J.R. Effect of acute hypoxia on the hormonal response to exercise. J. Appl.
Physiol. 42:587-592, 1977.
28. Sutton, J.R., and L. Lazarus. Effects of adrenergic blocking agents on growth hormone
responses to physical exercise. Hormone Metab. Res. 6:428-429, 1974.
29. Wagenmakers, A.J.M. Role of amino acids and ammonia in mechanisms of fatigue. In
Muscle Fatigue Mechanisms in Exercise and Training (Med. Sport Sci. Vol. 34), P.
Marconnet,P.V. Komi, B. Saltin, and0.M. Sejersted(Eds.). Basel, Switzerland:Karger,
1992, pp. 69-86.
30. Wolfe, R.R. Protein-Energy Interactions (Proceedings of an International Dietary
Energy Consultancy Group). Geneva: Nestle Foundation, 1991.
31. Young, A.J., W.J. Evans, A. Cymerman, K.B. Pandolf, J.J. Knapik, and J.T. Maher.
Sparing effect of chronic high altitude exposure on muscle glycogen utilization during
exercise. J. Appl. Physiol. 525357462, 1982.
32. Young, P.M., J.R. Sutton, H.J. Green, J.T. Reeves, P.B. Rock, C.S. Houston, and A.
Cymerman. Operation Everest 11: Metabolic and hormonal responses to incremental
exercise to exhaustion. J. Appl. Physiol. 73:2574-2579, 1992.

Acknowledgments
This work was supportedby the French Ministry of Research and Technology(Project
Aliment 2000). Dietary supplements were generously provided by Dietetique & Sante S.A.,
B.P. 106, Revel, France.

Manuscript received: February 6, 1995


Acceptedforpublication: April 18, 1996

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