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Acute Hormonal and Neuromuscular Responses

and Recovery to Forced vs. Maximum
Repetitions Multiple Resistance...

Article in International Journal of Sports Medicine · August 2003

DOI: 10.1055/s-2003-41171 · Source: PubMed

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 Acute Hormonal and Neuromuscular Responses and J. P. Ahtiainen1
A. Pakarinen2
Recovery to Forced vs. Maximum Repetitions Multiple W. J. Kraemer3
Resistance Exercises K. Häkkinen1
Physiology & Biochemistry

Abstract
Acute hormonal and neuromuscular responses and recovery
three days after the exercises were examined during the maxi-
mum repetitions (MR) and forced repetitions (FR) resistance ex-
ercise protocols in 16 male athletes. MR included 4 sets of leg
and GH (p < 0.01) were larger in FR than in MR. The decrease of
56.5 % (p < 0.001) in maximal isometric force in FR was greater
(p < 0.001) than that of 38.3 % in MR (p < 0.001) and force re-
mained lower (p < 0.01) during the recovery in FR compared to
MR. The larger decrease in isometric strength in FR than in MR
was also associated with the decreased maximal voluntary EMG

Downloaded by: University of Jyvaskyla. Copyrighted material.

presses, 2 sets of squats and 2 sets of knee extensions (with
12 RM) with a 2-min recovery between the sets and 4 min be-
tween the exercises. In FR the initial load was chosen to be higher
than in MR so that the subject could not lift 12 repetitions per set
by himself. After each set to failure the subject was assisted to
perform the remaining repetitions to complete the 12 repetitions
per set. Thus the exercise intensity was greater in FR than in MR.
Both loading protocols led to the great acute increases (p < 0.05 ±
0.001) in serum testosterone, free testosterone, cortisol and GH
concentrations. However, the responses in cortisol (p < 0.05)
410
of the loaded muscles. The data indicate that the forced repeti-
tion exercise system induced greater acute hormonal and neuro-
muscular responses than a traditional maximum repetition exer-
cise system and therefore it may be used to manipulate acute re-
sistance exercise variables in athletes.
Key words
Anabolic hormones ´ isometric force ´ EMG ´ neuromuscular fa-
tigue ´ muscle damage ´ bodybuilding

Introduction heavy-resistance exercise when performed repeatedly (i. e. train-

Heavy resistance exercise is a potent stimulus for acute increases
in the concentrations of circulating hormones in young men.
These acute increases are highly dependent on the type of resist-
ance exercise protocol, i. e. number of sets and repetitions per set,
length of rest period between sets and muscle mass involved em-
ployed [e. g. 16, 23].
Especially anabolic hormones (e. g. growth hormone, testoster-
one) influence growth and development [10]. The stress of
ing) is a stimulus for both strength development and muscle fi-
ber hypertrophy. This may be related, at least in part, to exercise-
induced acute increase in endogenous anabolic hormones [23].
The hormonal response to resistance exercise is also associated
with changes in the neuromuscular performance. Intense mus-
cular work used typically during a heavy-resistance training
leads to a momentary decrease in strength accompanied by de-
creases in voluntary maximal neural activation of the loaded
muscles [14,17, 21]. The effect of the fatiguing load on the neuro-

Affiliation
1
Neuromuscular Research Center & Department of Biology of Physical Activity, University of Jyväskylä,
Jyväskylä, Finland
2
Department of Clinical Chemistry, University of Oulu, Oulu, Finland
3
The Human Performance Laboratory, Department of Kinesiology, The University of Connecticut,
Storrs, CT, USA

Correspondence
J. Ahtiainen ´ Department of Biology of Physical Activity ´ University of Jyväskylä ´ P.O. Box 35 ´
40014 University of Jyväskylä ´ Finland ´
Phone: +358-14±2602083 ´ Fax: +358-14±2602071 ´ E-Mail: ahtiainen@sport.jyu.fi

Accepted after revision: November 20, 2002

Bibliography
Int J Sports Med 2003; 24: 410±418  Georg Thieme Verlag Stuttgart ´ New York ´ ISSN 0172-4622
muscular system is related not only to the intensity of exercise
but also to the specific type of the fatiguing load, and the recov-
ery time between high-intensity intermittent contractions [e. g.
17, 25]. It is speculated that there might be ªan optimalº level of
acute neuromuscular fatigue after each strength training session
to stimulate adaptation processes. According to the principle of
progressive strength training, the next training session should
usually take place under the conditions where complete recov-
ery has taken place. Therefore, the magnitude of temporary neu-
romuscular fatigue and the rate of recovery after intensive
heavy-resistance exercise may be an indication of its effective-
ness for long-term adaptations of the neuromuscular system. Re-
covery also involves the coordinated functioning of several phys-
iological processes (i. e. regeneration of disrupted muscle fibers)
that are heavily influenced by the availability and actions of
specific hormones [29, 31, 34].
Traditionally in strength training various exercises have been
performed using a so-called maximum repetition system (i. e.
each set is performed to a momentary concentric failure). In or-
der to overload the muscle progressively, the training intensity
and/or volume and/or frequency should be increased periodical-
ly. Because of the risk for overtraining it is appropriate for long-
term strength training purposes to prioritise the increase in the
training intensity. In practical training the term ªintensityª has
been used to define the magnitude of the load employed or the
rate of work performed [3]. In strength training the intensity
can also be modified by special training systems, such as so
called ªforced repetitionsº as defined by Fleck and Kraemer [9].
Forced repetitions are a special resistance training system, which
strength athletes, especially bodybuilders, use to increase train-
ing intensity. Forced repetitions means that, after the trainee has
achieved a momentary concentric failure (i. e. a set until exhaus-
tion has been performed), a training partner will assist by lifting
or pushing the load just enough to allow the trainee to complete
three to four additional repetitions. This system ªforcesº the
trainee to continue to produce force, although he or she is al-
ready extremely fatigued. It is suggested that it is necessary to
achieve the failure, i. e. the momentary muscular fatigue during
the resistance exercise sets to gain maximally muscle mass and
strength. Although it might be reasonable to suggest that the
forced repetition training system may be an effective way to
stimulate muscles, no previous studies have been conducted on
acute, short- or long-term hormonal and/or neuromuscular re-
sponses of the forced repetitions training system.
This study examined the influence of the exercise intensity to
acute hormonal and neuromuscular responses and short-term
recovery. For that purpose the responses of the forced repetitions
training system were compared to those produced by the tradi-
tional maximum repetitions training protocol. The experimental
training session was created in both loading conditions so that it
included multiple exercises, leg press, squat and knee extension,
as usually used in training for the leg extensor muscles in ath-
letes like bodybuilders.
Ahtiainen JP et al. Hormonal Responses to Resistance Exercise ¼ Int J Sports Med 2003; 24: 410 ± 418
Material and Methods
Subjects
Sixteen healthy physically active men volunteered to participate
in this investigation (mean  SD; age = 26.8  3.5 y, height =
180  6 cm, weight = 81  9 kg, body fat = 13.8  3.5 %). Each sub-
ject had recreational experience with resistance training but
none were competitive strength athletes. No medication was
taken by the subjects, which would have been expected to affect
physical performance. Each subject was informed of the poten-
tial risks and discomforts associated with the investigation and
all the subjects gave their written informed consent to partici-
pate. The Ethics Committee of the University of Jyväskylä
approved the study.
Physiology & Biochemistry
Experimental design and loading protocols
The subjects were familiarized with the experimental testing
procedures on the control day about 1 week before the measure-
ments. Anthropometrical measurements and resistance load ver-
ifications for each experimental exercise were also determined at
this time. The percentage of body fat was estimated from meas-
urements of skinfold thickness [6].
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During the control day three blood samples were obtained from
each subject. One blood sample was drawn in the morning after
twelve hours of fasting and approximately eight hours of sleep
for the determination of basal serum hormone concentrations.
Two blood samples were also drawn within 1/2 h without exercise
at the same time of day that each subject would later undertake
his heavy resistance loading protocols to determination of nor-
mal diurnal variation of serum hormone concentrations.
The experimental design comprised two loading sessions sep- 411
arated by two weeks performed at the same time of day (Fig. 1).
Recovery of the loading sessions was examined for three days
after both loadings. The first one of the loading sessions was a
so-called maximum repetition (MR) protocol. MR included 4
sets of leg press (David 210, from a knee angle of 59.5  2.08 to
1808 = knee straight), 2 sets of squats (Smith machine, from a
knee angle of 708 to 1808) and 2 sets of knee extensions (David
200, from a knee angle of 59.4  1.68 to 1808) with a two-minute
recovery between the sets and four minutes between the exerci-
ses. All the sets were performed with the maximum load possi-
ble for 12 repetitions (12 RM). The second loading session was a
so-called forced repetition (FR) protocol. In FR the loading proto-
col was the same as in MR, but the initial load was set approxi-
mately 15 % higher than in MR so that the subject could not per-
form twelve repetitions without assistance and would require
assistance during the last three to five repetitions. The foot posi-
tions and exercise machine settings were identical in both load-
ings. In FR the exact force of the assistance was measured. In the
squat the assistant and the subject stand on the separate force
plates. The force plates were used to measure the accurate force
of the assistance given by the assistant while ªliftingº the barbell
with his hands during the concentric phase of the squat. In the
leg press and knee extension exercises the handle force dyna-
mometer was attached to the foot support and was used to
measure the accurate force given by the assistant during his
ªpullingº action while the subject was extending the knees. The
assistant was the same person in all measurements. The external

Physiology & Biochemistry
412
force produced by the assistant during the concentric phases of
the exercises was analysed and then subtracted from the total
volume of the work (loads ” sets ” reps) to determine actual total
work performed by the subject in the FR loading. The deepness of
the squats was controlled by a sound signal. The duration of the
concentric phases of dynamic muscle actions was measured by
an electronic goniometer placed on the knee joint.
Neuromuscular measurements
An electromechanical dynamometer was used to measure maxi-
mal voluntary isometric force of the bilateral leg extension ac-
tion at a knee angle of 1078 before, after two and four sets of leg
press, after squats and after knee extensions as well as 24 h, 48 h
and 72 h after the loadings. Electromyographic activity (EMG)
was recorded from the agonist muscles vastus lateralis (VL) and
vastus medialis (VM) of the right leg during the maximal isomet-
ric action. Bipolar surface electrodes (Beckman miniature-sized
Ahtiainen JP et al. Hormonal Responses to Resistance Exercise ¼ Int J Sports Med 2003; 24: 410 ± 418
Fig. 1 Experimental design
of the study.
Downloaded by: University of Jyvaskyla. Copyrighted material.
skin electrodes 650 437, Illinois, USA) with 20 mm interelectrode
distance were employed. The electrodes were placed longitudi-
nally over the muscle belly on the motor point area determined
by an electrical stimulator (Neuroton 626). The positions of the
electrodes were marked on the skin by small ink dots to ensure
the same electrode positioning in each test during the experi-
mental period [15]. EMG signals were recorded telemetrically
(Glonner Biomes 2000) and stored on magnetic tape (Racall 16)
and to the computer with CODAS computer system (Dataq In-
struments, Inc.). EMG signal was amplified (by a multiplication
factor of 200, low-pass cut-off frequency of 360 Hz 3dB±1) and
digitized at a sampling frequency of 1000 Hz. EMG was full-
wave rectified, integrated (iEMG in mV ” s) and time normalized.
The activity (iEMG) of the VL and VM was averaged and analysed
in the maximal force phase (500 ± 1500 ms) of the isometric
muscle actions [17].
Blood collection and analyses
Blood samples were drawn from the antecubital vein via multi-
ple venipunctures for the determination of serum total and free
testosterone, cortisol and growth hormone concentrations. Fin-
gertip blood samples were drawn for the determination of blood
lactate. Hemoglobin and hematocrit were also determined to es-
timate changes in plasma volume (PV). During the loading ses-
sion blood samples were drawn before, after two and four sets
of leg press, after squats and after knee extensions as well as 15
and 30 min after the loadings. In the morning of the first and sec-
ond day after the loadings, fasting blood samples were obtained
for the determination of basal hormone concentrations. Venous
blood samples were drawn for the determination of serum crea-
tine kinase (CK) activity before as well as 24, 48 and 72 h after
the loadings at the same time of day that each subject had done
his heavy resistance loading protocols. All blood samples were
obtained at the same body position of the subject.
Serum samples for the hormonal analyses were kept frozen at
±20 8C until assayed. Serum testosterone concentrations were
measured by the Chiron Diagnostics ACS:180 automated chemi-
luminescene system using ACS:180 analyzer (Medfield, MA). The
sensitivity of the testosterone assay was 0.42 nmol/l, and the in-
tra-assay coefficient of variation was 6.7 %. The concentration of
serum free testosterone was measured by radioimmunoassays
using kits from Diagnostic Products Corp. (Los Angeles, CA). The
sensitivity of the free testosterone assay was 0.52 pmol/l, and the
intra-assay coefficient of variation was 3.8 %. The assays of serum
cortisol were carried out by radioimmunoassays using kits from
Farmos Diagnostica (Turku, Finland). The sensitivity of the corti-
sol assay was 0.05 nmol/l, and the intra-assay coefficient of var-
iation was 4.0 %. Concentrations of growth hormone were meas-
ured using radioimmunoassay kits from Pharmacia Diagnostics
(Uppsala, Sweden). The sensitivity of the GH assay was 0.2 g/l,
and the intra-assay coefficient of variation was 2.5 ± 5 %. All the
assays were carried out according to the instructions of the man-
ufacturers. All samples for each test subject were analysed in the
same assay for each hormone [15]. Hormone concentrations
were not corrected for plasma volume changes since the target
tissues sense the actual molar concentrations [22]. Blood lactate
concentrations were determined using a Lactate kit (Roche, Ger-
many). Serum CK activity was determined using a Creatine Ki-
nase kit (Roche, Germany).
Recovery after the loadings
The rate of recovery after the MR and FR loadings was studied at
24 h, 48 h and 72 h after the loadings. The measurements were
done at the corresponding time of the day as the subject's heavy
resistance loading protocols. The recordings of maximal isomet-
ric force and concurrent EMG evaluated the recovery of the neu-
romuscular performance after the loadings. Serum CK activity,
subjective muscle soreness (DOMS) and muscle swelling were
also determined as markers of muscle disruption possibly caused
by the resistance exercises. DOMS was rated on a scale of 0 ( = no
pain) to 10 ( = maximum pain) for the overall muscle soreness of
the quadriceps muscles. To examine muscle swelling, the thick-
ness of the vastus lateralis was measured at the middle of the
thigh with ultrasound (US) (Aloka SSD-2801s). The US measure-
ments were taken twice at each time point and the mean of the
two measurements was used in the statistical analysis. Acute
Ahtiainen JP et al. Hormonal Responses to Resistance Exercise ¼ Int J Sports Med 2003; 24: 410 ± 418
Fig. 2 The loads in FR vs. MR sets (mean  SE). *Significantly different
(** = p < 0.01, *** = p < 0.001) from set 1 value of each exercise. #Sta-
Physiology & Biochemistry
tistically significant difference (# = p < 0.05, ## = p < 0.01) between the
MR vs. FR loadings.
muscle swelling was also measured by examining effects of the
resistance exercise on the thickness of the loaded muscles. Blood
samples for the determination of basal hormone concentrations
were drawn from each subject after twelve hours of fasting and
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approximately eight hours of sleep in the first and second morn-
ings after the loadings.
Statistical analyses
Standard statistical methods were used for the calculation of
means, standard deviations (SD), standard errors (SE) and Pear-
son bivariate correlation coefficients. The changes in the vari-
ables over time from the pre-level were analysed using general
linear model (GLM) analysis of variance with repeated measures.
Differences between the variables within each time point were
analysed utilizing paired±samples of t-tests. The p < 0.05 criteri- 413
on was used for establishing statistical significance.
Results
Loading
According to the design of the study, the mean ( SD) load was
13.2  11.2 % higher in all FR sets than in MR sets (p < 0.001)
(Fig. 2). The average duration of the concentric phases was
28.4 % longer in FR (1537  467 ms) than in MR (1100  257 ms)
(p < 0.001). In general, assistance was given for 6.0  3.2 repeti-
tions of the 12 repetitions sets in the FR loading. The total vol-
ume of the work (loads ” sets ” reps) in MR was 12 616  1807 kg
and in FR 14126  1744 kg (p < 0.001). However, when the
amount of external force produced by the assistant during the
concentric phases of the FR repetitions was taken into account,
the actual total volume of FR did not differ from that of MR.
Acute neuromuscular responses
Isometric force
Significant decreases (p < 0.05 ± 001) occurred in maximal iso-
metric force in both loading protocols (Fig. 3). Maximal isometric
force decreased during the entire course of the loading in MR
down to 61.7  9.3 % (from 3024  463 N to 1859  383 N,
p < 0.001) and in FR down to 43.5  12.2 % (from 3031  498 N to
1290  360 N, p < 0.001). The decrease in isometric force was
greater (p < 0.01 ± 0.001) in FR than in MR during the entire
Physiology & Biochemistry
Fig. 3 Maximal voluntary isometric leg extension force and maximal
integrated EMG activity (mean  SE) during and after the MR vs. FR
loadings (% from pre-loading value). *Significantly different
(* = p < 0.05, ** = p < 0.01, *** = p < 0.001) from corresponding pre-
exercise value. # Statistically significant difference (# = p < 0.05,
##
= p < 0.01, ### = p < 0.001) between the MR vs. FR loadings.
course of the loadings. The decreased isometric force remained
lower (p < 0.01) in FR than in MR for 24 h (Fig. 3). In FR maximal
isometric force was still lower (p < 0.05) at the third day after the
loading as compared to the pre-level.
414
EMG activity
There were no significant changes in the maximum integrated
EMG of the isometric action at any time points in MR, but FR led
to the decreases (p < 0.05 ± 0.01) in the EMG (Fig. 3). The changes
in maximal isometric force and the changes in EMG correlated
with each other after the entire FR loading (r = 0.51, p < 0.05).
The EMG was significantly lower (p < 0.05 ± 0.01) in FR compared
to MR after the second and fourth leg press sets and after the
squat sets. There were no statistically significant alterations in
EMG activity during the recovery as compared to the pre-level
or between the loadings.
Blood lactate
The blood lactate concentration increased up to 14.2  3.2 mmol/l
(p < 0.001) and to 15.0  2.8 mmol/l (p < 0.001) after the entire
course of the MR and FR loadings, respectively. There were no
significant differences between the loadings.
Plasma volume
PV decreased after the entire loading by ±11.8  6.8 % (p < 0.001)
and ±14.4  3.3 % (p < 0.001) in MR and FR, respectively, with no
significant differences between the loadings.
Ahtiainen JP et al. Hormonal Responses to Resistance Exercise ¼ Int J Sports Med 2003; 24: 410 ± 418
Fig. 4 Serum testosterone and free testosterone concentrations
(mean  SE) during the MR vs. FR loadings. *Significantly different
Downloaded by: University of Jyvaskyla. Copyrighted material.
(* = p < 0.05 ** = p < 0.01 *** = p < 0.001) from corresponding pre-ex-
ercise value. #Statistically significant difference (# = p < 0.05) between
the MR vs. FR loadings.
Acute hormonal responses
Control samples
There were no significant differences in the concentrations of se-
rum hormones examined between the two control blood sam-
ples drawn during the control day with no exercise (data not
shown).
Exercise samples
Serum testosterone concentrations increased after the entire
course of the MR and FR loadings from 22.5  7.0 nmol/l up to
25.2  9.2 nmol/l (p < 0.05) and from 23.2  7.9 nmol/l up to
26.8  7.5 nmol/l (p < 0.05), respectively (Fig. 4). After the MR
loading serum testosterone concentrations decreased signifi-
cantly below the pre-level (p < 0.05, post 30 min).
Serum free testosterone concentrations increased after the entire
course of the MR and FR loadings from 53.8  13.9 pmol/l up to
72.5  27.9 pmol/l (p < 0.001) and from 58.3  18.1 pmol/l up to
79.1  25.7 pmol/l (p < 0.01), respectively (Fig. 4). After the MR
loading serum testosterone concentrations decreased signifi-
cantly below the pre-level (p < 0.05, post 30 min). There were no
significant differences between the MR and FR loadings.
Serum cortisol concentrations increased after the entire course
of the MR and FR loadings from 0.36  0.12 mol/l up to
0.53  0.13 mol/l (p < 0.001) and from 0.35  0.11 mol/l up to
0.65  0.11 mol/l (p < 0.001), respectively (Fig. 5). The acute re-
sponses in cortisol (p < 0.05) were larger in FR than in MR after
the squat and knee extension sets as well as 15 and 30 minutes
after the loadings. The changes in maximal isometric force and
the changes in serum cortisol concentrations correlated with
each other after the entire FR loading (r = -0.55, p < 0.05).
Fig .5 Serum cortisol concentrations (mean  SE) during the MR vs. Fig. 6 The relative changes in serum GH concentrations (meanSE)
FR loadings. *Significantly different (* = p < 0.05, ** = p < 0.01,

Physiology & Biochemistry

*** = p < 0.001) from corresponding pre-exercise value. ##Statistically
significant difference (p < 0.01) between the MR vs. FR loadings.
during the MR vs. FR loadings. *Significantly different (* = p < 0.05,
** = p < 0.01, *** = p < 0.001) from corresponding pre-exercise value.
#
Statistically significant difference (# = p < 0.05, ## = p < 0.01) between
the MR vs. FR loadings.

Serum GH concentrations increased from 1.0  1.9 mg/l up to

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Table 1 Basal hormone values (mean  SD) of the first and second
23.6  15.2 mg/l (p < 0.001) and from 0.3  0.8 mg/l up to
mornings after the loadings
28.6  16.3 g/l (p < 0.001) after the entire course of the MR and
FR loadings, respectively. The relative changes in GH concentra- Control day 1st morning 2nd morning
tions were greater (p < 0.05 ± 0.01) in FR than in MR already after morning after the after the
(±1 week) loading loading
the four sets of leg press, and remained larger throughout the en-
tire loading and the recovery of 30 min (Fig. 6). The changes in Testosterone (nmol/l) 25.5  7.2 MR 24.8  7.1 25.6  6.8
blood lactate and the changes in serum GH concentrations cor- FR 25.9  8.0 25.0  6.7
related with each other in both MR (r = 0.66 , p < 0.01) and FR Free testosterone (pmol/l) 60.1  16.8 MR 70.3  26.9 67.3  17.9**
(r = 0.55 , p < 0.05) loadings after the two sets of knee extensions. FR 65.1  20.1 69.6  20.6*
Cortisol (mol/l) 0.46  0.14 MR 0.46  0.14 0.46  0.14
CK-activity and muscle soreness FR 0.50  0.14 0.45  0.10 415
CK-activity peaked at 24 hours after MR and FR from
128.4  114.8 IU/l up to 237.8  127.7 IU/l (p < 0.01) and from * p < 0.05; ** p < 0.01 compared to the control day

67.9  55.7 IU/l up to 251.8  168.4 IU/l (p < 0.001), respectively.

Subjective muscle soreness ratings peaked at 48 hours after MR
and FR from 0.2  0.3 up to 3.4  2.0 (p < 0.001) and from 0.2  0.3
up to 3.7  2.9 (p < 0.001), respectively.

Thickness of the vastus lateralis
VL thickness increased after the entire course of the loadings by
9.3  5.1 % (p < 0.001) and by 11.8  5.7 % (p < 0.001) in MR and FR,
respectively. The values decreased gradually during the recovery
period in both loadings.
Basal hormone concentrations
Basal hormone concentrations were unaltered compared to the
control day morning values after the loadings (Table 1), except
for the increased free testosterone values at 48 hours after the
MR (p < 0.01) and FR (p < 0.05) loading protocols.
Discussion
The primary findings of this study were that both maximum rep-
etition and forced repetition loading protocols led to the great
acute hormonal responses. There were no differences between
the loadings in the acute testosterone and free testosterone re-
sponses, while the cortisol and relative GH responses were great-
er in FR than in MR. Both loading protocols led also to major neu-
romuscular fatigue observable with the acute decreases in iso-
metric force associated with high blood lactate concentration,
while only FR led to the decrease in the maximal voluntary EMG
of the loaded muscles. Also the recovery in maximal isometric
force after the loading was slower in FR than MR.
The present study was, to our knowledge, the first one to exam-
ine the exercise and recovery profiles of acute hormonal respon-
ses when the exercise regimen included several multiset exerci-
ses for the same muscle group (i. e. leg press, squat, and knee ex-
tensions for the quadriceps femoris) as used in typical hyper-
trophic type of strength training. Forced repetitions are a special
resistance training system, which is used to increase the intensi-
ty of training. It may not be common among strength trainers to
perform forced repetitions in every set and more than three to
four of them per set, but in the present study the number of the
forced repetitions was intentionally high in order to create the
different experimental conditions between the two loadings.
The MR loading was performed first in the experimental design

Ahtiainen JP et al. Hormonal Responses to Resistance Exercise ¼ Int J Sports Med 2003; 24: 410 ± 418
 due to practical reasons. In this way it was possible to obtain the
true strength level of the subjects at the moment and to create
maximal loading for both loading protocols. It cannot be exclud-
ed, but there are no serious reasons to believe that the acute hor-
monal or neuromuscular responses could be affected by the pre-
vious loadings in the present study. However, contrary to acute
hormonal and neuromuscular responses the muscle damage
markers as CK-activity, subjective muscle soreness and muscle
swelling could be diminished on the second experimental load-
ings via prophylactic effect of the previous loading [28].
Testosterone promotes muscle hypertrophy via enhancing pro-
tein synthesis and may also contribute to force production by
its potent influence on neural mechanism (e. g. increased neuro-
Physiology & Biochemistry
transmitter synthesis) [4, 27, 29, 30]. Heavy resistance exercise-
induced acute increases in serum testosterone concentration
may be caused by the influence of the increased circulation in
the testicles, activation of the sympatic nervous system, in-
creased lactate accumulation and/or luteinizing hormone con-
centrations [7, 8,18]. Changes in the plasma volume and the in-
creased clearance of the circulating testosterone may also, at
least in part, explain the exercise-induced testosterone response.
In the present study serum testosterone and free testosterone
concentrations increased significantly during both loading pro-
tocols with no significant differences between the loadings. The
contribution of free testosterone represents the amount of bioac-
tive testosterone, which can act directly with androgen receptors
in the target tissue (e. g. skeletal muscle) to mediate changes of
the function of a muscle cell via enhanced protein synthesis. Re-
gardless of the mechanism(s) of an exercise-induced increase of
serum testosterone concentrations, the skeletal muscle will be
exposed to an elevated peripheral testosterone concentration
416 and thus the likelihood of possible interactions with potential
muscle cell receptors could increase. Furthermore, the increase
of testosterone concentration in serum has been connected to
up-regulation of androgen receptors [5]. In the present study,
serum testosterone concentrations decreased 30 minutes after
MR below the pre-level, which is probably caused by the lowered
LH response of the testicles due to the exercise-induced increase
in serum testosterone concentration. Interestingly, serum total
testosterone concentration turned to a downward trend already
after the squat exercise, especially in FR. This may be due to
smaller activated muscle mass in the knee extension exercise,
lowered blood lactate concentration, failure in testosterone pro-
duction and/or increase in its hepatic clearance. Contrary to total
testosterone, serum free testosterone increased throughout the
entire loading. The concentration of serum free testosterone
was also increased on the second morning after both loadings as
compared to the control day value. That is perhaps an indication
of a compensation mechanism of the hormonal system against
the exercise-induced stress.
Cortisol influences, among others, the metabolism of amino
acids and glucose. Cortisol increases gluconeogenesis and recy-
cling of proteins. Amino acid availability is an important regula-
tor of the muscle protein metabolism [2]. In the postexercise re-
covery period cortisol contributes to maintain sufficient rates of
glycogen synthesis, protein turnover and supply of protein syn-
thesis by amino acids [33]. On the other hand, the exercise-in-
duced increase in cortisol concentration may inhibit the secre-
Ahtiainen JP et al. Hormonal Responses to Resistance Exercise ¼ Int J Sports Med 2003; 24: 410 ± 418
tion of GnRH [26] and/or the increased level of ACTH may com-
pete with LH of the androgen receptors of Leydig cells [1]. In the
present study the exercise-induced cortisol response was signif-
icantly greater in FR than in MR. The exercise response occurred
in FR after the four sets of leg press and in MR after the squat,
when the threshold for the cortisol response was probably ex-
ceeded [32]. Exercise-induced cortisol response may be due to
glycolytic demands of the exercise, stimulated effect of catechol-
amines and/or a consequence of neural control of muscle work.
The exercise response of endocrine action is triggered by the cen-
tral motor command and the responses are further supported by
positive feedback influences from proprio- and metaboreceptors
in muscles [20]. In the present study, the changes in maximal
isometric force and changes in EMG as well as the changes in
maximal isometric force and the changes in serum cortisol con-
centration correlated significantly between each other in FR. This
may partly support the hypothesis for the influence of neural
control of the muscle work to exercise-induced cortisol response.
GH has anabolic effects on the muscle cell by increasing trans-
portation of amino acids in the muscle cell and increasing pro-
tein synthesis. GH is an anabolic hormone, and therefore, heavy
Downloaded by: University of Jyvaskyla. Copyrighted material.
resistance exercise-induced increased secretion of GH may be
important for the process of training-induced muscle hypertro-
phy [24]. In the present study, serum concentrations of GH in-
creased greatly after both loadings but the relative change in
the GH response was significantly greater in FR than in MR. Exer-
cise-induced increase in serum GH concentrations measured by
the immunoreactive method (molecular size 22kDa) can be ex-
plained mainly by hypoglycemia and the stimulatory effect of
the motor cortex and the symphatic nervous system of the hypo-
thalamus. Further explanation might also be an increased acidity
in the muscle caused by anaerobic muscle work, which stimul-
ates metaboreceptors and sends afferent feedback to the central
nervous system and hypothalamus leading to an increased secre-
tion of GH [11,12,19]. This is supported by the present study
showing that serum GH concentrations correlated with blood
lactate concentrations in both loadings. In the present study, the
deceased maximal voluntary isometric actions were also asso-
ciated with the decreased EMG activity of the loaded muscles
during the FR loading. Therefore greater relative increase in GH
concentrations in FR may indicate the importance of the central
motor command to the exercise-induced GH response.
The investigation of the rate of recovery after the heavy resist-
ance exercise may be advantageous to estimate a proper strength
training frequency and/or intensity and/or volume. Especially
the forced repetitions resistance exercise system is supposed to
cause muscular soreness [9]. Mechanical load due to resistance
exercise may cause some level of myofibrillar disruptions to the
activated muscles. In the present study maximal isometric force,
subjective muscle soreness, muscle swelling and CK activity
were used to estimate the level of muscle damage caused by the
heavy resistance exercise and rate of recovery after the loadings.
Maximal isometric force was significantly more lowered in FR
than in MR during the recovery and isometric force remained
significantly lowered even on the third day after FR. There were
no significant alterations in the maximum iEMG during the re-
covery period. Thickness of the vastus lateralis (i. e. muscle swel-
ling) increased during the present loading session and returned
to the pre-exercise level on the next day. Although the loading
regimens were strenuous, the muscle damage markers as CK-
activity and subjective muscle soreness did not increase greatly
after the present kinds of high load and low velocity types of dy-
namic muscle actions.
The load was greater in FR especially at the beginning of the
loading session than in MR. However, when the force of the
assistance was reduced from the total load, there was no signifi-
cant difference between FR and MR in the total work that the
subjects had performed by themselves. Nevertheless, FR led to a
greater decrease in maximal isometric force than MR. That may
be due to a greater accumulation of lactic acid to working mus-
cles during FR and/or the decrease in the ability to activate motor
units (especially type II) by the central nervous system [17]. The
larger increase in blood lactate concentration and the greater de-
crease in EMG in FR compared to MR supports this suggestion. It
has been demonstrated earlier that the decrease in maximal
strength during high-intensity fatiguing strength training ses-
sions may be associated with the decrease in the maximal volun-
tary neural activation of the exercised muscles [14,17]. The pres-
ent FR protocol may produce greater peripheral and central fa-
tigue compared to MR where fatigue was caused mostly by the
peripheral factors. The greater blood lactate concentration may
be due to a longer concentric phase of the dynamic muscle ac-
tions in FR than in MR as a consequence of the higher load used
in FR. Therefore, blood circulation may decrease in the activated
muscles and acid metabolic waste products accumulated in the
muscle cells [13]. The larger decrease in isometric force associa-
ted with decreased neural activation of the loaded muscles and
the higher GH response in FR than in MR indicate that the usage
of the forced repetitions protocol may be beneficial for the devel-
opment of muscle mass and muscle strength during strength
training. On the other hand, the lowered recovery of force pro-
duction in FR may indicate an increased risk for overtraining if
the training frequency is kept too high and/or the overall volume
of each training session is too high.
In summary, the present data showed that FR compared to MR
loading led to greater stress for the neuromuscular system
shown through the larger decrease in isometric force and iEMG
of the loaded muscles. Furthermore, the recovery of the isomet-
ric force was slower after the FR than MR loading. Because the
degree of acute neuromuscular fatigue and the time needed for
recovery may differ considerably between the MR and FR loading
protocols, there is a need to optimize the contents and the fre-
quency of different training sessions in order to create proper
strength training programs to match the individual require-
ments of athletes. The FR loading stimulated the hormonal sys-
tem, especially the secretion of GH, more than MR did. Whether
the repeated usage of the multiple FR protocol is associated with
larger gains in muscle mass or strength during prolonged
strength training, and how any differential gains are related to
acute and long term endogenous anabolic adaptations needs to
be examined.
Ahtiainen JP et al. Hormonal Responses to Resistance Exercise ¼ Int J Sports Med 2003; 24: 410 ± 418
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