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maximal exercise
GAITANOS, G. C., C. WILLIAMS, L. H. BOOBIS, AND S. subsequent bouts of exercise (10, 31). However, these
BROOKS. Human muscle metabolism during *intermittent maxi- suggestions were on the basis of blood lactate measure-
mal exercise. J. Appl. Physiol. 75(2): 712-719, 1993.-Eight ments only. Other studies using brief work periods (lo-
male subjects volunteered to take part in this study. The exer- 15 s) interrupted by short intervals of rest (E-40 s) have
cise protocol consisted of ten 6-s maximal sprints with 30 s of suggested that the contribution of glycogenolysis to the
recovery between each sprint on a cycle ergometer. Needle total energy demand was either of minor importance (26)
The average l-s integral of effective load and flywheel they were all verbally encouraged throughout each 6-s
speed, over the total time period of each exercise period sprint. After the last sprint the subjects rested for 10 min
(i.e., 6 s), is referred to as mean power output (MPO). on an examination couch adjacent to the cycle er-
Each sprint was initiated from a stationary start because gometer.
of the increased accuracy and ease with which the start- On a separate day, maximum oxygen uptake (VO, max)
ing point could be identified by computer. was determined using a graded cycling test. The first
Subjects. Eight healthy male physical education stu- work load was high enough to ensure that exhaustion
dents, whose mean age, height, and weight were 26.7 t would occur within 6-12 min, the load being increased by
8.4 (SD) yr, 175.6 t 11.4 cm, and 71.8 t 11.4 kg, respec- 0.5 kg at the end of every 3 min. Expired air samples were
tively, gave their informed consent and volunteered to collected during the final minute of each load increment
take part in this study, which was approved by the Uni- and at the last 60 s of exercise. Heart rate was monitored
versity Ethical Committee. continuously by cardiac oscillos,cope during all the exer-
Protocol. All subjects were familiarized with sprint cy- cise tests and was visible to the experimenters through-
cling until fully confident of producing an all-out effort out the tests.
from a stationary start. They were instructed to refrain Blood samples. Venous blood samples were taken from
from any form of intense physical exercise on the day an antecubital vein by using an intravenous cannula (16
before testing. On the day of the experiment, subjects gauge, Venflon) fitted with an injection diaphragm. The
reported to the laboratory after a fast of at least 4 h. They cannula was inserted under local anesthesia (0.5 ml of
6 8
Sprint No FIG. 1. Schematic i llustration of study
protocol. ret, recovery; post, postexercise.
Muscle
sampling
t
Blood
sampling
714 MUSCLE METABOLISM DURING MAXIMAL EXERCISE
n Peak Power(W)
Mean POwer(w)
1 2 3 4 5 6 7 8 9 10
Sprint No.
FIG. 2. Peak and mean power output for each of ten 6-s sprints. FIG. 3. Blood lactate concentrations during exercise and recovery.
Values are means + SD; n = 7. Values are means + SD; n = 7 except as indicated. Pre, preexercise;
Post 1 and 5, after sprints 1 and 5, respectively; Pre 10 and Post 10,
before and after sprint 10, respectively; 3, 5, and 10 Post, 3,5, and 10
norepinephrine by using high-performance liquid chro- min after last sprint, respectively.
matography with electrochemical detection (3).
Muscle samples. Muscle biopsy samples were taken ables. Statistical significance was accepted at the 5%
under local anesthesia (2 ml, 1% lidocaine) at rest before level. Results are presented as means + SD.
warm-up and immediately after the first sprint from one
leg and 10 s before and immediately after the last sprint
TABLE 1. Muscle metabolites before and after first and last 6-s sprint of multiple sprint test on cycle ergometer
Metabolites Pre-1st 6 s (n = 8) Post-1st 6 s (n = 8) Pre-10th 6 s (n = 7) Post-10th 6 s (n = 7)
to be zero (range O-23.1 mmol ATP/kg dry wt, n = 7). mmol l kg dry wt- l s- 2.3kO.6 0.3+0.5*
Glycogenolytic and glycolytic rates. The minimum an- Rate of glycogen degradation,
aerobic glycogenolytic and glycolytic rates were esti- mmol glucosyl units. kg dry wt- l s- 7.2t4.1 3.4k3.0"
mated using the formulas described by Hultman and Sjo- Values are means + SD; n, no. of subjects. * Significant difference
holm (12). as follows: glvcogenolvsis = AG-6-P + (0.33 X from szwint I. P < 0.01.
MUSCLE METABOLISM DURING MAXIMAL EXERCISE 717
ten 6-s sprints. These decrements in power output were be attributed, in part, to a high rate and great transfor-
accompanied by a considerable reduction in the rate of mation of inactive phosphorylase b to active phosphory-
ATP production by anaerobic processes. This was lase a resulting from the rapid fivefold rise in plasma
mainly reflected by the lo-fold decrease in the rate at epinephrine concentration that was observed after the
which glycogen was degraded to lactate in the final sprint first sprint.
compared with the values calculated for the first sprint. In the final 6 s of exercise the estimated ATP produc-
Previous studies from this laboratory have proposed tion rate from anaerobic sources was reduced to 35.6%
that intermittent maximal exercise of brief duration (6 s) (5.3 t 2.5 mmol . kg dry wt- s-l) of that estimated for
l
with short rest intervals (30 s) leads to an increasing the first sprint. Glycolysis contributed only 16.1% as the
demand on anaerobic glycolysis to maintain the rate of glycolytic rate had fallen 7.6 times. In addition there was
energy production because of incomplete resynthesis of a lo-fold decrease in the glycogenolytic rate, although it
PCr stores (31). This was not the case in the present always exceeded glycolysis.
study, which employed a similar work-to-rest ratio. The increase in muscle lactate concentration found at
The initial ATP production rate, estimated from the the end of the exercise test is associated with large reduc-
changes in the concentration of muscle metabolites, was tions in muscle pH (11). Blood pH was also decreased to
14.9 t 2.2 mmol kg dry wt-l
l ls-l. This value is close to 7.10 units after completion of the test. This blood pH
the theoretical maximum rate of 17 mmol. kg dry value has been related to decreases in muscle pH to -6.6
wt-l l s-l calculated by McGilvery (21). Degradation of (11). By using the equation described by Sahlin (24) for
PCr accounted for one-half of the ATP resynthesized calculating the muscle pH from lactate and pyruvate
lated more lactate in muscle (r = 0.79, P < 0.05; n = 8) ported by Harris and co-workers (8). On the basis of
and also had the greatest ATP production rates from these calculations it appears that the muscle PCr stores
anaerobic sources (r = 0.84, P < 0.01; n = 8), supporting were reduced almost to the same extent during the first
the view that most of the glycogen was metabolized an- and the final sprints. If this were the case, then it sug-
aerobically. These findings suggest that the MPO gener- gests that a 30-s recovery interval may be sufficient to
ated over the first sprint must have been fueled by energy allow significant PCr resynthesis and, therefore, consid-
derived mainly from PCr and anaerobic glycogenolysis. erable contribution from PCr to ATP production during
These results also support the observations of other au- this type of intermittent exercise, even after ten maximal
thors (2,14) that glycogenolytic processes leading to lac- 6-s sprints. This is in contrast to previous suggestions
tate formation are initiated within the first few seconds that PCr degradation may be of minor importance in
of maximal dynamic exercise. maintaining high rates of ATP production during the
The extent to which glycogen was used anaerobically latter stages of intermittent exercise of maximal inten-
in the first 6-s bout of exercise, as calculated from the sity (31). Therefore, the decline in power output in the
accumulation of hexose monophosphates, lactate, andpy- present study was accompanied by evidence of decreas-
ruvate, represents one of the highest rates of glycogenoly- ing contribution to ATP production from glycogen degra-
sis (4.4 t 0.9 mmol glucosyl units kg dry wt?. s-l) re-
l dation to lactate formation as indicated by the changes in
corded for human quadriceps femoris muscle. This high muscle hexose monophosphates and lactate concentra-
value possibly reflects the great involvement of the vas- tions before and after the last exercise bout. However,
tus lateralis muscle group during maximal cycling. This whether the decreased rate of ATP resynthesis in the last
value is similar to that observed in a previous study from sprint, as a result of a low rate of glycogen degradation to
this laboratory (2). lactate, was secondary to inhibited ATP hydrolysis, as
The high glycogenolytic rate calculated from changes previously suggested (l3), is not known.
in metabolites during the first sprint requires a total Despite a dramatic reduction in energy yield from an-
phosphorylase activity of 264 mmol glycosyl units kg l aerobic glycogenolysis in the last sprint, the MPO
dry wt- ls-. This greater phosphorylase activity could achieved was 73% of that in the initial sprint. Several
718 MUSCLE METABOLISM DURING MAXIMAL EXERCISE
possibilities exist to explain this difference. First, a output generated by the subjects is being measured dur-
greater reliance on aerobic metabolism of glycogen is pos- ing this type of treadmill sprinting (17).
sible. A detectable fall in glycogen content was observed In summary, the results of the present study indicate
in the last bout of exercise, which cannot be accounted that the average power output over the first sprint was
for by the accumulation of glycolytic intermediates, glu- supported by energy that was derived mainly from PCr
cose, or lactate. It is difficult to account for the amount of and anaerobic glycogenolysis. In the final sprint, how-
lactate that escaped from the muscle into the circulation ever, no lactate accumulation was apparent, yet the aver-
during that sprint. The increase in blood lactate immedi- age power output was still 73% of that in the initial
ately after exercise may reflect glycogenolytic activity of sprint. On the basis of the circumstantial evidence that
all preceeding exercise bouts. Because no differences in suggests an inhibition of anaerobic glycogenolysis at the
blood lactate were seen before and after the last exercise end of exercise, it is reasonable to propose that ATP re-
bout, however, it is possible that little lactate left the synthesis was mainly derived from PCr degradation and
muscle during this period of exercise. Moreover, the post- oxidative metabolism.
exercise hyperglycemia in blood observed in the present
study is thought to have been derived from liver glycogen- Address reprint requests to C. Williams.
olysis due to the high levels of circulating catechol- Received 19 July 1991; accepted in final form 11 March 1993.
amines.
The possibility, therefore, that some aerobic metabo- REFERENCES
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