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The impact of metabolic stress on hormonal


responses and muscular adaptations
ARTICLE in MEDICINE &AMP SCIENCE IN SPORTS &AMP EXERCISE JUNE 2005
Impact Factor: 4.46 DOI: 10.1249/01.mss.0000170470.98084.39 Source: PubMed

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The Impact of Metabolic Stress on Hormonal


Responses and Muscular Adaptations
KAZUSHIGE GOTO1, NAOKATA ISHII2, TOMOHIRO KIZUKA1, and KAORU TAKAMATSU1
Institute of Health and Sport Sciences, University of Tsukuba, Tsukuba, Ibaraki, JAPAN; and 2Department of Life
Sciences, Graduate School of Arts and Sciences, University of Tokyo, Komaba, Tokyo, JAPAN
1

ABSTRACT
GOTO, K., N. ISHII, T. KIZUKA, and K. TAKAMATSU. The Impact of Metabolic Stress on Hormonal Responses and Muscular
Adaptations. Med. Sci. Sports Exerc., Vol. 37, No. 6, pp. 955963, 2005. Purpose: The purpose of this study was to examine the impact
of exercise-induced metabolic stress on hormonal responses and chronic muscular adaptations. Methods: We compared the acute and
long-term effects of an NR regimen (no-rest regimen) and those of a WR regimen (regimen with rest period within a set).
Twenty-six male subjects were assigned to either the NR (N 9), WR (N 9), or control (CON, N 8) groups. The NR regimen
consisted of 35 sets of 10 repetitions at 10-repetition maximum (RM) with an interset rest period of 1 min (lat pulldown, shoulder
press, and bilateral knee extension). In the WR regimen, subjects completed the same protocol as the NR regimen, but took a 30-s rest
period at the midpoint of each set of exercises in order to reduce exercise-induced metabolic stress. Acute hormonal responses to both
regimens were measured followed by a 12-wk period of resistance training. Results: Measurements of blood lactate and serum hormone
concentrations after the NR and WR regimens showed that the NR regimen induced strong lactate, growth hormone (GH), epinephrine
(E), and norepinephrine (NE) responses, whereas the WR regimen did not. Both regimens failed to cause significant changes in
testosterone. After 12 wk of resistance training, the NR regimen caused greater increases in 1RM (P 0.01), maximal isometric
strength (P 0.05), and muscular endurance (P 0.05) with knee extension than the WR regimen. The NR group showed a marked
increase (P 0.01) in muscle cross-sectional area, whereas the WR and CON groups did not. Conclusion: These results suggest that
exercise-induced metabolic stress is associated with acute GH, E, and NE responses and chronic muscular adaptations following
resistance training. Key Words: MUSCULAR STRENGTH, MUSCULAR HYPERTROPHY, REST PERIOD, LACTATE

exercise intensity (10RM) and shorter rest periods (1 min)


considerably elevate GH and TES concentrations, whereas
those using higher intensity (5RM) and a longer rest period
(3 min) do not. The former protocol is known as a hypertrophy-type regimen, and is used typically by bodybuilders.
It has been known that local accumulation of metabolic
subproducts such as lactate and hydrogen ions stimulates
sympathetic nerve activity and exercise-induced catecholamine secretion (3). The similar afferent pathway has been
shown recently to play an important role in the regulation of
anabolic hormone secretions from the hypothalamus-pituitary (25,29). For example, Gordon et al. (6) have demonstrated the effects of alkalosis treatment by sodium bicarbonate (NaHCO3) ingestion on exercise-induced hormonal
response. The NaHCO3 treatment resulted in a greater pH
and attenuated GH response to 90 s of cycle ergometer
sprinting than the placebo treatment. Moreover, exercise
with vascular occlusion markedly enhances metabolite accumulation within the muscles and concomitant GH secretion (25,29), but not TES (29). Although the mechanism by
which anabolic hormone secretions, especially GH, are
stimulated by acid base changes is yet to be fully understood, it is possible that activation of the hypothalamus
pituitary axis by afferent signals from muscle metaboreceptors has a role in this action (6,25,28). Hence, an exercise
regimen with greater metabolic stress should cause greater
anabolic hormone and catecholamine responses to resistance exercise.

t has been shown that resistance training for several


weeks increases muscle cross-sectional area, strength,
and power (2,24). Development of optimal resistance
exercise regimens has been a major focus of physiologists
and trainers. Muscular hypertrophy and strength gains following resistance training are thought to be dependent on
the intensity of exercise, in such a way that an intensity of
more than 65% of 1-repetition maximum (1RM) is required
to achieve a substantial effect (16).
The mechanism(s) underlying these muscular adaptations
involves many factors, that is, mechanical, metabolic, endocrine and neural factors. Of these factors, training-induced muscular hypertrophy might be at least partially related to the secretions of endogenous anabolic hormones
such as growth hormone (GH) and testosterone (TES) (5).
The type of exercise regimen has a significant affect on the
magnitude of anabolic hormonal responses (4,7,23). Kraemer et al. (13) have shown that regimens using moderate

Address for correspondence: K. Takamatsu, Ph.D., Institute of Health and


Sport Sciences, University of Tsukuba, 1-1-1 Tennodai, Tsukuba, Ibaraki,
305-8574, Japan; email: ktaka@taiiku.tsukuba.ac.jp.
Submitted for publication August 2004.
Accepted for publication December 2004.
0195-9131/05/3706-0955/0
MEDICINE & SCIENCE IN SPORTS & EXERCISE
Copyright 2005 by the American College of Sports Medicine
DOI: 10.1249/01.mss.0000170470.98084.39

955

In addition, it has been suggested that metabolic stress has


a substantial role in stimulating muscular hypertrophy and
strength gains (20,21,22,26). Schott et al. (21) have reported
that an isometric training regimen resulting in a greater
decrease in intramuscular pH induced a greater muscle
hypertrophy compared with a regimen with only small
change in pH. Rooney et al. (20) revealed that improvement
in strength following 6 wk of dynamic resistance training
was significantly greater in the regimen without rest compared with a regimen with a 30-s rest period between each
repetition, although the magnitude of muscular hypertrophy
was not evaluated. The enhanced metabolic accumulation
within muscles caused by vascular occlusion resulted in
marked increases in muscular strength (22) and muscle
cross-sectional area (26) following resistance training.
These findings suggest that exercise-induced metabolic
changes may be associated with muscular adaptations to
resistance training. Moreover, these effects might be induced, in part, by secretion of anabolic hormones, but no
previous studies have been conducted on the influences of
exercise-induced metabolic changes on acute hormonal responses and long-term adaptations of muscles.
Therefore, the purpose of the present study was to examine the impacts of exercise-induced metabolic stress on
acute hormonal responses and chronic muscular adaptations
following a 12-wk training period. For this purpose, we
compared the acute and long-term effects of two exercise
regimens with same relative intensity and volume, that is, a
regimen without rest period within a set (no-rest regimen)
and a regimen with a rest period (rest regimen).With continuous repetitions, the former exercise regimen would
cause the marked change in acid base balance. Conversely,
brief rest periods inserted into the middle of each set would
reduce accumulation of metabolites, thereby resulting in an
attenuation of exercise-induced metabolic changes (21). We
hypothesized that a regimen with intraset rest period would
cause smaller anabolic hormone and catecholamine responses, and also smaller increases in muscular size and
strength after the period of training.

METHODS
General Procedure
Twenty-six healthy male subjects (age, 22.7 0.5 yr;
height, 172.0 1.1 cm; body mass, 65.9 1.5 kg) participated in this study. They were undergraduate or graduate
students at the University of Tsukuba, and all had experience with recreational resistance training. However, none of
the subjects were involved in any regular training program
at the beginning of the study. No medication (e.g., anabolic
steroids, creatine, sympathoadrenal drugs) was taken by the
subjects, which would have been expected to affect physical
performance. They were informed about the experimental
procedure as well as the purpose of the present study, and
their written informed consent was obtained in advance. The
study was approved by the ethics committee for human
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Official Journal of the American College of Sports Medicine

experiments at the Institute of Health and Sport Sciences of


the University of Tsukuba.
The subjects were assigned to experimental (N 18) and
untrained control (CON; N 8) groups on the basis of
muscular strength and physical characteristics measured before the training period. The experimental group was further
divided into a no-rest regimen (NR) group (N 9) and
regimen with rest period (WR) group (N 9). Subjects
were matched according to physical characteristics, training
history, and muscular strength, and there were no significant
differences for any parameter between the groups.
The present study consisted of two separate experiments.
In the first part of the experiment, acute hormonal responses
to the NR and WR regimens were investigated using the
subjects in the NR group. In the second part of the experiment, the long-term effects of 12 wk of resistance training
with NR and WR regimens were investigated. Muscular
strength, endurance, and the cross-sectional area (CSA) of
the quadriceps femoris (QF) were measured before and after
the period of training.
Study of Acute Hormonal Response
Subjects and exercise protocol. Nine male subjects
in the NR group participated in this experiment. They performed three exercises for the upper and lower limbs (lat
pulldown, shoulder press, and bilateral knee extension) using weight-stack machines. These three exercises were chosen for this study because they represent upper- and lowerlimb exercises commonly used in exercise programs.
Although several studies have used whole-body programs
with higher work volume (13,15), our study included only
three upper- and lower-limb exercises, for the reason that
the subjects were unable to adhere strictly to a whole-body
exercise regimen with a greater work volume and metabolic
stress at the beginning of the training period. Furthermore,
a low-volume regimen using four exercises (bench press, lat
pulldown, knee extension, leg curl) has been shown to
produce significant elevations of lactate and GH (14).
The NR regimen (regimen without rest) consisted of 10
repetitions 35 sets (three sets for the lat pulldown and
shoulder press, and five sets for the knee extension) at
10RM. The subjects were allowed to rest for 1 min between
all the sets and exercises. In the first set of exercise, the
exercise intensity was set at approximately 75% of 1RM
(10RM). Thereafter, the intensity was adjusted to allow the
subjects to complete 10 repetitions in each set. In the WR
regimen, the subjects completed the same protocol as the
NR regimen, but took a 30-s rest period at the midpoint of
each exercise set (between the fifth and sixth repetitions).
The subjects performed the exercise at the same relative
intensity, number of repetitions in each set, and interset rest
period as those for the NR regimen. The WR regimen with
the intraset rest period was designed to reduce the accumulation of metabolites in the muscles without changing the
major variables of the exercise regimen (i.e., total contraction time, work volume, number of sets, rest period between
sets). The acute experiment involving NR and WR regimens
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TABLE 1. Physical characteristics of the subjects.


Variable
Age (yr)
Pre
Height (cm)
Pre
Body mass (kg)
Pre
Post
% fat
Pre
Post

NR group
(N 9)

WR group
(N 9)

CON group
(N 8)

23.1 0.9

21.9 0.7

23.3 0.9

171.7 2.3

170.3 1.7

174.3 1.2

65.2 3.0
66.4 2.8**

66.4 2.4
66.9 2.3

66.3 2.2
66.8 2.4

20.3 1.2
18.8 1.3*

20.9 1.4
20.2 1.4

19.6 1.2
20.1 1.3

Values are means SE. Pre and post indicate the values obtained before and after the
training period, respectively. * Significant difference from pretraining value (P 0.05);
** significant difference from pretraining value (P 0.01).

was separated by more than 5 d. The subjects were instructed to lift and lower the load at a constant velocity,
taking about 2 s for each concentric and eccentric action. All
exercise sessions were preceded by stretching of the major
muscle groups and a single set of warm-up exercises at 50%
of 1RM.
Blood sampling and analyses. Following an overnight fast, the subjects came to the laboratory at 9:30 am and
rested for 30 min before the first blood collection. Venous
blood samples were obtained from an indwelling cannula in
the antecubital vein before, and at 0 (immediately after the
exercise), 15, and 30 min after exercise. Serum samples for
hormone analyses were stored frozen at 85C until analysis. Serum GH concentration was measured by radioimmunoassay (RIA) using kits from Daiichi Radioisotope Lab,
Japan. The sensitivity of the GH assay was 0.05 ngmL1,
and the inter- and intraassay coefficients of variation (CV)
were 3.6 and 3.4%, respectively. Serum testosterone (TES)
concentration was measured by RIA using kits from DPC
Corporation, Japan. The sensitivity of the TES assay was 5.0
ngdL1, and the inter- and intraassay CV were 8.4 and
5.9%, respectively. Plasma concentrations of epinephrine
(E) and norepinephrine (NE) were measured by high-performance liquid chromatography (HPLC) using kits from
Tosoh Corporation, Japan. The sensitivity of these assays,
and inter- and intraassay CVs were 6.0 pgmL1, 2.7, and
2.0% for E; and 6.0 pgmL1, 2.4, and 1.3% for NE. Blood
samples were also obtained from the fingertip to measure
lactate concentration (LA) using an automatic lactate analyzer (YSI1500 Sport, Yellow Springs Instruments, U.S.).
Long-Term Effects of Resistance Training
Subjects and exercise protocol. Twenty-six male
subjects were assigned to the CON, NR, and WR groups.
The NR and WR groups performed the same exercise regimens as those used in the experiments on acute hormone
responses (see above). The physical characteristics of the
subjects are shown in Table 1. Resistance training using lat
pulldown, shoulder press and bilateral knee extension was
performed for 12 wk (23 sessions in total). Exercise training
was performed only once in the first week, and thereafter
twice per week until the 12th week. The training sessions
involved one-on-one supervision by the same assistant
ROLE OF METABOLITES IN RESISTANCE TRAINING

trainer throughout the experimental period. The subjects


were asked to refrain from performing other resistance training during the experiment period.
Measurements of muscular strength. Muscular
strength was evaluated as 1RM for the shoulder press and
knee extension exercises. Maximal isometric and isokinetic
strengths were also measured for the knee extension exercise. Before measuring 1RM, a warm-up with 10 repetitions
at 50% of the perceived 1RM and stretching of the major
muscle groups subjected to the exercises were performed. In
the present study, 1RM was basically measured by progressively increasing the load until the subjects were unable to
perform a lift. However, when the 1RM exceeded the
equipped weight of the machines, an estimation method
using submaximal load was applied. The relationship between RM and %RM was as follows: 2RM 95% of 1RM;
3RM 92.5% of 1RM; 4RM 90% of 1RM. Before this
study, the reliability of this estimation was confirmed by
comparing the actual and estimated 1RM in the same group
of subjects. This comparison showed that the intraclass
correlation coefficient was greater than 0.90, and confirmed
reliability was high.
Maximal isometric and isokinetic strengths of the unilateral knee extension exercise were measured using an isokinetic dynamometer (COMBIT, Minato Medical Science, Japan). The subjects were familiarized with the test procedure on
several occasions before taking measurements. They sat on a
chair with their right leg (dominant side) attached firmly to the
lever of the dynamometer. Maximal isometric strength was
measured at a knee angle maintained at 100 (180 at full
extension). The subjects were instructed to exert maximal force
for 3 s. The highest value of 23 trials was adopted. Maximal
isokinetic strength at 60, 120, 180, 240, and 300 s1 was then
measured. Three to five repetitions were carried out to determine the peak torque for joint angle ranging between 90 and
180. A rest period of 2 min was allowed between each trial.
The order of angular velocities tested was randomized, and the
same order was used in each subject between before and after
training period.
Evaluations of muscular endurance. Before and
after the period of training, muscular endurance of the upper
and lower limbs was evaluated by the maximal number of
repetitions for the shoulder press and bilateral knee extension exercises with the same relative load, that is, 70% of
pre- or posttraining 1RM. Each subject was instructed to
perform the exercise at a frequency of 30 repetitions per
minute until exhaustion. Muscular exhaustion was defined
as the moment when the weight ceased to move or the
subjects failed to maintain the prescribed pace. Any repetition without full range of movement was not counted. Following the test, the performed exercise volume was calculated (load repetition) and used as a measure of muscular
endurance.
Measurements of muscle cross-sectional area.
The CSA of the thigh at its midpoint was measured using
nuclear magnetic resonance imaging (MRI) with a body coil
(MRI; 1.0 T, Magnetom Impact, Siemens, U.S.). Fifteen
serial sections with a sectional thickness of 15 mm were
Medicine & Science in Sports & Exercise

957

acquired with the field of view, repetition time, and echo


time being 240 mm, 800 ms, and 20 ms, respectively, and
the scan matrix 256 256. Image acquisition was started
after the subject lied in the supine position with his legs
extended and relaxed. The scan time was approximately
1315 min.
From the obtained serial sections, those for two portions
near the midpoint (halfway between the trochanter major
and head of the tibia) of the thigh were chosen for the
analysis of muscle CSA. On the selected cross-sectional
images, the outlines of quadriceps femoris (QF) were traced
by the same expert. Traced images were inputted into a
computer (Power Macintosh G4, Apple Computer), and the
CSA of the QF were measured using NIH image software
(version 1.61). The maximal isometric strength per unit of
CSA was then determined as an index of neuromuscular
function. The measurements were repeated twice for each
image, and the mean values were adopted. A strong correlation between the first and second measurements (r 0.99)
indicated a high reliability of the measurements.
Statistical analysis. Data are expressed as means
SE. In the study on acute hormonal response to the NR and
WR regimens, a two-way analysis of variance (ANOVA)
with repeated measures was used. In the event of a significant F-ratio, a Tukeys HSD post hoc test was used to
compare means. In the study on the long-term effects of
resistance training, a two-way ANOVA with repeated measures and a Tukeys HSD post hoc were used. Differences
between the percent changes among the three (NR, WR,
CON) groups were examined by a one-way ANOVA followed by a Tukeys HSD post hoc test. A selective bivariate
relationship was investigated using a Pearson productmoment correlation coefficient. P 0.05 was considered as
significant.

RESULTS
Acute hormonal and lactate responses. Figures 1
and 2 show acute changes in blood LA, serum GH, and TES
(Fig. 1), and plasma E and NE (Fig. 2) before the period of
training. No significant difference was seen in resting LA
and hormone concentrations between the exercise regimens.
In the NR regimen, LA and hormone concentrations (except
for that of TES) showed significant elevations after exercise,
whereas those of the WR regimen showed no significant
changes except for LA and NE. Between the NR and WR
regimens, significant differences were observed in the postexercise concentrations of LA, GH, and NE, among which
GH showed the largest difference. When the GH response was
assessed by the area under the time-concentration relationship (GHauc), the value after the NR regimen (559.6
200.9 ngmL1) was approximately threefold higher than that
after the WR regimen (185.0 86.7 ngmL1; P 0.05).
However, no significant difference was seen in TES concentrations between the two regimens, although the postexercise
value (30 min) in the NR regimen was lower than its preexercise value (P 0.05).
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Official Journal of the American College of Sports Medicine

FIGURE 1Acute changes in blood lactate, growth hormone, and


testosterone concentrations after exercises with the NR and WR regimens before the period of training. Values are means SE (N 9).
* Significant difference from preexercise value (P < 0.05); # significant
difference between the regimens (P < 0.05).

Changes in muscle cross-sectional area and


body composition. Changes in physical characteristics
over the period of training are shown in Table 1. No significant difference was observed in the pretraining values
between the groups. For the NR group only, body mass
http://www.acsm-msse.org

FIGURE 3Percent changes in muscle cross-sectional area (CSA)


after exercise training in the no-rest regimen (NR; N 9), regimen
with a rest period within a set (WR; N 9), and untrained (CON; N
8) groups. Values are means SE; ** significant difference between
the groups (P < 0.01).

FIGURE 2Acute changes in epinephrine and norepinephrine concentrations after exercises with the NR and WR regimens before the
period of training. Values are means SE (N 9). * Significant
difference from preexercise value (P < 0.05); # significant difference
between the regimens (P < 0.05).

significantly increased, and the percentage of fat significantly decreased after a 12-wk training period, indicating an
increase in lean body mass (LBM).
Figure 3 shows the percent changes in muscle CSA of QF
after the 12-wk training period. To reduce errors in the
measurement associated with a slight mismatch between the
sectional portions obtained before and after the training
period and incidental deformations of muscles during the
MRI processes, two sections around the mid portion of the
thigh were selected, and the mean tissue CSA was obtained
from these sections. No significant difference was observed
in the pretraining values of CSA between the groups. The
CSA in both the NR (12.9 1.3%) and WR (4.0 1.2%)
groups significantly increased after the training period, and
the change in CSA was significantly larger in the NR group
than in the WR group (P 0.01). No significant change in
CSA was seen in the CON group after the training period
(0.3 1.3%, NS).
The percent change in CSA after the 12-wk training
period did not show clear correlations with acute exerciseROLE OF METABOLITES IN RESISTANCE TRAINING

induced GH increase (difference from preexercise: r 0.66,


P 0.05) and GHauc (r 0.66, P 0.05). However, a
slightly clearer correlation was found between the peak
value of GH after acute exercise and the percent change in
CSA following the training period (r 0.67, P 0.05).
Hormones other than GH did not show a positive correlation
with the percent change in CSA.
Changes in muscular strength. Changes in 1RM
during and after the training period are shown in Table 2 and
Figure 4. No significant differences were observed in the
pretraining values between the groups. In both the NR and
WR groups, 1RM of the shoulder press and knee extension
exercises significantly increased after the training period,
and posttraining increases were also significantly greater in
the NR and WR groups than in the CON group. However,
no significant difference in 1RM of the shoulder press was
seen between the effects of the NR and WR regimens. On
the other hand, 1RM of the knee extension showed a significantly larger increase in the NR group (66.4 5.2%)
than in the WR group (39.0 3.7%, P 0.01).
TABLE 2. Changes in muscular strength after the training period.
Variable

NR group

WR group

CON group

Maximal isometric strength (Nm)


Pre
251.0 15.2
266.6 18.4
247.4 16.9
Post
300.6 23.0** 282.7 16.8
251.1 17.1
1RM (kg)
Shoulder press
Pre
60.3 3.9
60.3 3.6
55.3 3.9
Post
77.9 4.6** 73.0 2.1** 54.0 3.3
Knee extension
Pre
69.3 4.2
71.9 3.8
78.5 5.3
Post
114.6 6.7** 99.3 4.5**
79.3 5.1
Values are means SE. Pre and post indicate the values obtained before and after the
training period, respectively. ** Significant difference from pretraining value (P 0.01);
significant difference from CON group (P 0.05); significant difference from CON
group (P 0.01).

Medicine & Science in Sports & Exercise

959

FIGURE 4 Changes in one-repetition maximum (1RM) of shoulder


press and bilateral knee extension exercises during and after a 12-wk
training period. Absolute values (left) and percent changes (right) are
shown. Values are means SE; ** significant difference from pretraining value (P < 0.01); # significant difference from CON group
(P < 0.05); ## significant difference from CON group (P < 0.01);
significant difference between the groups (P < 0.01).

The changes in forcevelocity relationships after the


training period are shown in Figure 5. All values of isometric and isokinetic torques were normalized to the pretraining
values of isometric torque. The NR group showed significant increases in isometric and isokinetic strengths at almost
all velocities examined, whereas no significant changes
were observed in the WR and CON groups. When compared
between groups, isometric strength showed a significantly
greater increase in the NR group (19.1 3.1%) than in the
WR (7.2 3.2%) and CON (1.5 1.0%) groups. The
maximal isometric strength per unit of CSA did not significantly change after the training period in both the NR (3.6
0.2 vs 3.8 0.2 Nmcm2, NS) and WR (3.6 0.2 vs
3.7 0.2 Nmcm2, NS) groups. In addition, no significant
difference was seen in the percent changes of maximal
isometric strength per unit of CSA between the groups,
suggesting that the strength gain in NR group was caused
primarily by muscular hypertrophy.
Changes in muscular endurance. Muscular endurance was evaluated as the exercise volume performed at
70% of 1RM for the upper- and lower-limb muscles (Table
3 and Fig. 6). In the upper-limb muscles, no significant
changes were seen in the NR and WR groups after the
training period. In the lower-limb muscles, muscular endurance in the NR and CON groups significantly improved
after training period, whereas that in the WR group did not.
In addition, the percent change in exercise volume was
significantly greater in the NR group (41.8 10.2%) than
in the WR (7.8 8.0%) and CON (5.9 1.7%) groups,
indicating that only the NR regimen was substantially effective in improving muscular endurance.
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Official Journal of the American College of Sports Medicine

FIGURE 5Changes in forcevelocity relationships after the period of


training. All values of knee extension strength were normalized to the
pretraining values of isometric strength (0s1). Values are means
SE; * significant difference from pretraining value (P < 0.05); ** significant difference from pretraining value (P < 0.01).

DISCUSSION
This study showed that a NR regimen caused larger
elevations of blood LA, GH, and NE concentrations than a
WR regimen. In addition, training with NR regimen caused
http://www.acsm-msse.org

TABLE 3. Changes in exercise volume during shoulder press and knee extension
exercises at 70% of 1RM after the training period.
Variable
Shoulder press (J)
Pre
Post
Knee extension (J)
Pre
Post

NR group

WR group

CON group

609.8 58.7
681.1 41.0

546.9 36.3
624.3 17.3

454.3 49.0
453.6 43.1

884.8 44.2
931.8 45.1

897.7 75.1
955.6 92.6*

878.6 71.3
1219.5 86.5**#

Values are means SE. Pre and post indicate the values obtained before and after the
training period, respectively. Exercise volume was calculated as load number of
repetitions. * Significant difference from pretraining value (P 0.05); ** significant
difference from pretraining value (P 0.01); # significant difference from WR group (P
0.05); significant difference from CON group (P 0.01).

much larger increases in muscle CSA and strength than with


the WR regimen, although both regimens had the same
relative intensity and volume. As these regimens were designed to induce different metabolic responses, the specific
adaptations in these regimens may be related to differences
in exercise-induced metabolic stress.
The effects of the NR regimen in inducing elevations of
GH, E, and NE concentrations are consistent with previous
studies (12,23). However, the secretions of GH and NE were
abolished when a short rest period (30 s) was inserted in the
middle of an exercise set (WR regimen). The attenuation of
hormone responses in the WR regimen may result from a
smaller amount of metabolite accumulation within the muscles, because the blood lactate concentration after the WR
regimen was significantly lower than after the NR regimen
(Fig. 1). Several studies have shown that accumulation of
metabolic subproducts stimulates the secretions of GH and
catecholamine through afferent signals from intramuscular
metaboreceptors (6,25,28). This is consistent with the previous studies that have shown that alkalosis treatment attenuates exercise-induced GH secretion (6), and that ischemic exercise with increased metabolite accumulation
causes greater GH and NE responses (25,29). Therefore, it
appears that intraset rest in the WR regimen reduces acidity
via lactate production, causing smaller GH and NE responses. However, activation of motor centers may also
directly stimulate the hypothalamuspituitary axis and sympathoadrenal secretion (11). A more fatiguing NR regimen
might cause a stronger activation of the motor center and
thereby larger hormonal responses. Moreover, the traditional measurements of GH concentration have focused only
on the main 22 kDa isoform, whereas recent studies have
shown that other isoforms of GH have specific responses to
acute exercise (19). Further studies are needed to examine
changes in various GH isoforms after resistance exercise.
We hypothesized that a NR regimen with greater metabolic stress would produce larger hormonal responses than
a WR regimen. However, this hypothesis was not supported
for TES, because both regimens failed to evoke a significant
TES response, suggesting that TES responses are not greatly
affected by exercise-induced metabolic changes. This is
partially consistent with finding that muscle ischemia
caused by vascular occlusion did not enhance the TES
response to acute exercise (29). TES secretion is also suppressed for several hours after a single bout of strenuous
ROLE OF METABOLITES IN RESISTANCE TRAINING

FIGURE 6 Percent changes in work volume during shoulder press


and bilateral knee extension exercises at 70% of 1RM after the period
of training in the NR, WR, and CON groups. Values are means SE;
* significant difference between the groups (P < 0.05).

resistance exercise (18). However, in the present study,


measurements after the period of training showed small but
significant elevations of postexercise TES concentration
(preexercise: 626.8 15.6 ngdL1 vs mean of postexercise: 687.9 21.9 ngdL1, P 0.01). These changes in
acute TES responses might play a significant role in training-induced muscular adaptations (2).
The percent changes in 1RM and maximal isometric
strength were significantly greater in the NR group than in the
WR group (Figs. 4 and 5). In addition, maximal isokinetic
strength improved at almost all angular velocities in the NR
group, but not in the WR and CON groups (Fig. 5). Rooney et
al. (20) have shown a greater increase in dynamic strength in
a regimen with repeated six repetitions without rest compared
with a regimen with a 30-s rest period between each repetition.
Similarly, Schott et al. (21) have demonstrated that gains in
strength resulting from isometric training with long, fatiguing
Medicine & Science in Sports & Exercise

961

contractions are considerably greater than with a training regimen using shorter, intermittent contractions. The current and
previous results clearly indicate that continuous repetition
without pause is an important factor for strength gains following resistance training.
The present NR regimen caused an increase of approximately 13% in muscle CSA, whereas the WR regimen had
no such effect (Fig. 3). Recent studies have suggested that
enhanced metabolic stress within the muscles may strongly
stimulate protein synthesis and concomitant muscle hypertrophy (21,26,27). For example, Schott et al. (21) have
shown that an isometric training regimen with a greater
decrease in intramuscular pH could induce a larger degree of
muscle hypertrophy than that with a smaller pH change.
Takarada et al. (26) have also shown that resistance exercise
with moderate vascular occlusion provokes a marked increase in muscle CSA, and that this effect is related in part
to the increase in muscle fiber recruitment by acidic intramuscular environment. During exercise with marked metabolic changes, additional motor unit recruitment would be
induced to keep a given level of force, as shown by the
elevated electrical activity of the muscles 17). In the present
study, although electrical activity was not measured, we
speculate that greater metabolic stress in the NR regimen
would have affected muscle fiber recruitment. This might be
responsible for the larger muscular hypertrophy in the NR
regimen.
As shown in Figure 1, significant elevations of postexercise GH concentration were seen only in the NR regimen.
Although the actual roles of circulating anabolic hormones
in muscle growth are still unclear, combinations of GH and
mechanical loading would activate anabolic processes in
skeletal muscle (15). The acute elevation of GH has been
suggested to play a more significant role in increasing
insulin-like growth factor-1 (IGF-1) mRNA in the muscle
than do its chronic changes (9). In addition, lines of evidence have indicated that GH and IGF-1 play crucial roles
in the growth, development, and maintenance of skeletal
muscle. In the present study, the magnitude of the acute GH
responses showed a positive correlation with relative increases in muscle CSA (P 0.04 0.05), implying that GH
might contribute to exercise-induced muscular hypertrophy.
However, actions of GH for muscular hypertrophy are not
direct, and might be mediated by locally produced growth
factors (5). More research should be conducted to determine
whether GH plays a substantial role in resistance training
induced muscular hypertrophy.

Elevations of resting hormone concentrations may also be


related to muscular adaptations to resistance training. Previous studies have shown positive correlations between
changes in resting TES concentration and both muscular
strength development (2,10) and hypertrophy (8). However,
the present resting concentrations of GH, TES, IGF-1, and
cortisol did not change after the period of exercise training
in the NR group (data not shown).
Interestingly, a different adaptation in muscular endurance was observed between the upper and lower limbs. In
the lower limbs, improvement in muscular endurance was
significantly greater in the NR group than in the WR and
CON groups, whereas that in the upper limb was not regimen-dependent (Fig. 6). The reason for this is unclear, but
previous studies have suggested that upper-limb muscles
have a greater trainability than lower-limb muscles that are
more involved in daily physical activities (1). Therefore, a
considerable improvement in muscular endurance of the
upper limb might take place even after the WR regimen with
intraset rest period. The interpretation of muscular endurance data for knee extension needs precaution, because the
work volume in the CON group increased after the period of
training, possibly because of some familiarization with the
testing. Despite this fact, however, the improvement of work
volume during knee extension was much larger in the NR
group than in the WR and CON groups.
In conclusion, the NR regimen, consisting of moderate
intensity and short interset rest periods, evoked strong LA,
GH, E, and NE responses, and was also effective in increasing muscular size and strength after a period of training. In
contrast, both acute and long-term effects were markedly
diminished when a brief rest period was inserted into each
set of exercise. These results suggested that resistance exercise-induced metabolic stress was associated with acute
GH, E, and NE responses and chronic muscular adaptations.
Moreover, these indicated that reducing rest periods to increase metabolic stress was an effective strategy for gaining
substantial effects of resistance exercise. Development of
resistance exercise regimens designed to produce muscular
hypertrophy should take this phenomenon into account.
The authors are grateful to the subjects who participated in this
study. We are also grateful to Dr. Kaneko for assisting with the MRI
measurements, and Dr. Kraemer for his constructive comments.
The study was supported by grants from the Ministry of Education, Science, Sports and Culture of Japan, and from the Research
Fellowships of the Japan Society for the Promotion of Science for
Young Scientists.

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