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ISSN 03621197, Human Physiology, 2013, Vol. 39, No. 5, pp. 511–523. © Pleiades Publishing, Inc., 2013.
Original Russian Text © O.L. Vinogradova, D.V. Popov, A.I. Netreba, D.V. Tsvirkun, N.S. Kurochkina, A.V. Bachinin, Ya.R. Bravyi, E.V. Lyubaeva, E.A. Lysenko, T.F. Miller,
A.S. Borovik, O.S. Tarasova, O.I. Orlov, 2013, published in Fiziologiya Cheloveka, 2013, Vol. 39, No. 5, pp. 71–85.

Optimization of Training: New Developments

in Safe Strength Training
O. L. Vinogradovaa,b, D. V. Popova,b, A. I. Netrebaa, D. V. Tsvirkuna, N. S. Kurochkinaa,b,
A. V. Bachinina, Ya. R. Bravyia, E. V. Lyubaevaa, E. A. Lysenkoa, T. F. Millera,
A. S. Borovika, O. S. Tarasovaa,b, and O. I. Orlova
a
Institute of Biomedical Problems, Russian Academy of Sciences, Moscow, 123007 Russia
b
Biological Faculty, Moscow State University, Moscow, 119992 Russia
Received April 3, 2013

Abstract—The hypertrophic effect of strength training is known to be due to mechanical and metabolic stim
uli. During exercises with the restricted blood supply of working muscles, i.e., under the conditions of inten
sified metabolic stress, the training effect may be achieved with much lower external loads (20% of one repe
tition maximum). The effects of 8 weeks of highintensity (80–85% of one repetition maximum) strength
training were compared to lowintensity (50% of one repetition maximum) training without relaxation. The
highintensity strength training resulted in higher increases in strength and size of the exercised muscles than
training without relaxation. During highintensity training, at the muscle cross section, an increase in the
area occupied by type II fibers prevails; while, during training without relaxation, an increase in the area
occupied by type I fibers prevails. An exercise session without relaxation leads to a more pronounced increase
in the secretion of the growth hormone, insulinlike growth factor1, and cortisol. The expression of gene
regulating myogenesis (Myostatin) is changed in different ways after a highintensity strength exercise session
and after an exercise session without relaxation. Lowintensity strength training (50% of one repetition max
imum) without relaxation is an efficient way for inducing increases of the strength and size of the trained mus
cles. This lowintensive type of training may be used in rehabilitation medicine, sports, and fitness.

Keywords: strength training, skeletal muscle, muscle fibers, anabolic hormones, gene expression
DOI: 10.1134/S0362119713050162

Healthenhancing physical activity is a potent natu blood pressure, which increases the load on the heart).
ral drugfree way to improve the state of the human In this respect, there is an obvious need for safe weight
body. Aerobic exercises, which have been intensively training modes, which do not cause undesirable
studied, are practiced to improve the cardiovascular and changes at the system level but provide the changes in
respiratory systems. However, training in the anaerobic the metabolic processes in working muscles sufficient to
mode (strength training) has a more pronounced effect activate the growth of the muscle cells, i.e., the activa
on the musculoskeletal system. Strength training is an tion of intracellular signaling pathways and increasing
effective method to prevent decreases in the volume of the expression of genesregulator of myogenesis. The
skeletal muscles and their strength, as well as to prevent solution to this problem should be based on a scientifi
regulatory disorders that are related to these decreases. cally substantiated system of creating specific loads on
To date, it has been proved that the use of strength train the muscular system in order to correct irregularities in
ing can slow the progression of agerelated osteopenia the functioning of both the muscle and systemic regula
and sarcopenia, as well as the metabolic syndrome and tory mechanisms.
the related obesity, hyperlipidemia, and hypertension; it In this study, a method of lowintensity strength
can also reduce the development of rheumatoid myop training is described. This method involves training
athy and other kinds of myopathy [1]. without full muscle relaxation. It allows the strength of
Widespread use of strength training in prevention the trained muscles to be increased compared to tradi
and rehabilitation is hampered by the absence of a sci tional strength training, and, at the same time, the
entific approach to the optimization of training loads. It method is injuryfree.
is obvious that excessively small training loads are inef The hypertrophic effect of strength exercises is
fective, whereas excessive muscular contractions, in related to the influence of both mechanical and meta
addition to the risk of injuries of the musculoskeletal bolic stimuli. It is well known that strength exercises
system, may cause pronounced changes in the func impair cell membranes, which is confirmed by the sig
tioning of the autonomic vicseral systems (e.g., high nificant increase in the activity of creatine kinase (CK)

511
512 VINOGRADOVA et al.

in the blood 15–20 h after highintensity strength train It was shown using nuclear magnetic resonance
ing [2]. Mechanical stress, which is transmitted to the spectroscopy in vivo that, during lowintensity exercises
extracellular matrix of the muscle cell, can influence the with a restricted blood flow (30–40% 1 RM), metabolic
expression of gene that regulate myogenesis, intracellu changes in the working muscle are comparable with the
lar signaling, and control the rate of protein synthesis in changes that occur during traditional highintensity
muscle cells [3]. Such changes should be expected, strength training [25]. The authors conclude that, in
firstly, in strength training using highintensity loads. order to achieve significant metabolic changes during
This approach to strength training is very widespread; exercises with restricted blood flow, the load should be
thus, hereinafter, it is called the traditional strength at least 30–40% of 1 RM. Note that, in the vast majority
training. The main disadvantage strength training is the of studies on the effects of strength training with
high risk of injury of the musculoskeletal system in con restricted blood flow, the loads were 20% of 1 RM.
nection with the use of high training loads (75–90% of These studies involving untrained people or elderly sub
one repetition maximum (1 RM)). jects have shown the efficiency of the training programs
However, it is well known that strength exercises are used to increase their strength and muscle mass [5, 17,
accompanied by the accumulation of metabolites in 20, 26, 27]. It is likely that exercises with such loads are
muscles. The energy supply of the working muscle is suitable for rehabilitation; however, we assume that
mainly due to anaerobic metabolism, including the these loads will be insufficient for the activation of pro
pronounced activation of glycolysis. Due to the degra tein synthesis in the muscles of physically active people.
dation of creatine phosphate, in the muscle, there is a Also note that there are few studies comparing the effi
marked increase in the content of creatine [4, 5], which ciency of exercises with a restricted blood flow and tra
is the factor that stimulates anabolic processes [6, 7]. In ditional strength training [17, 27].
muscles and blood, there is a significant increase in the In this paper, we consider various modes for using
content of lactate, which, in turn, stimulates the secre the method of strength training without muscle relax
tion of anabolic hormones [8–10]. There is an increase ation with loads that are 50–70% of 1 RM, as well as
in the content of the somatotropic hormone, testoster regulatory changes resulting from this training, both at
one, and insulinlike growth factor 1 in the blood [11– the level of the whole body and within the working mus
14]. These metabolic changes may alter signaling and cles, including the changes in the expression of the
the expression of genes regulating myogenesis in muscle generegulating myogenesis. In addition, we present a
fibers [15–17]; and they may be related to an increase in comparison of the effects of strength training without
the muscle mass and strength [5]. Based on these con muscle relaxation and traditional highintensity
siderations, there were a number of attempts to potenti strength training.
ate the effects of strength training using exercises under
the conditions of the restricted blood supply of the
working muscles, when metabolic changes are intensi MATERIALS AND METHODS
fied, and, hence, the training effect could be achieved We compared the effects of 8 to 10week traditional
by using lighter external loads. Limiting the blood sup strength training (80–85% of 1 RM) and training with
ply was achieved with the help of occlusive cuffs [18– out relaxation with various loads (50–70% 1 RM) in leg
20] or by creating high pressure on the lower half of the and knee extensions (L series and K series) on the
body when a subject was placed in the pressure chamber strength and morphological characteristics of the work
[10, 21, 22]. However, the practical use of this approach ing muscles, as well as on the biochemical parameters of
is related to significant inconvenience to the subjects. blood and the hormonal status. We also studied the
Therefore, attempts were made to achieve similar effect of a single training session described above on the
effects by using more convenient and accessible expression of genes regulating myogenesis.
approaches. L series. Young, physically active male subjects
It is well known that, during the static tension, blood trained their leg extensor muscles (leg extension,
flow in a working muscle is reduced, and sometimes L series) for 8 weeks. Nine subjects from the control
completely absent; i.e., the conditions are similar to group (LC group) trained according to the traditional
occlusion. It is well known that exercises in a static (iso strength training routine; the remaining nine subjects
metric) mode are not very efficient for increasing the constituted an experimental group (LE group) and per
dynamic strength [23]. However, during the traditional formed lowintensity strength training without relax
strength training, continuous restriction of the local ation. The developing training for the LC group
blood flow cannot be achieved. This contradiction was included seven sets with 10min rest periods between
successfully resolved by developing a hybrid method of them. During each set, the training with a load of 80–
strength training, including static and dynamic compo 85% of 1 RM was performed to exhaustion (6–12 reps).
nents [24]. The method is based on the absence of relax The training movements (leg extension) were per
ation of the working muscles in the final phase of the formed with the greatest speed possible for them. The
training movement. This mode of training is called the time ratio of muscle tension and relaxation in this group
lowintensity method of strength training without was about 50% to 50%. For the LE group, the training
relaxation. session included 4 supersets for 3 sets, each with a load

HUMAN PHYSIOLOGY Vol. 39 No. 5 2013

 OPTIMIZATION OF TRAINING: NEW DEVELOPMENTS 513

of 50% of 1 RM. The duration of the set was 50–60 s, 40–50 s; the rest intervals between them were 30 s; and
the rest intervals between them were 30 s, and the inter the rest periods between the series were 5 min.
vals between the series were 10 min. The training move As can be seen from the above, the workload for the
ment was slower approximately by 3.5 times than during E group was increased in comparison to the series
the traditional strength training, at a constant speed. In described previously due to the greater intensity of exer
this group, continuous maintenance of muscle tension cising (65–70% vs. 50% of 1 RM in the previous exper
was achieved because the subjects did not carry the iment), and an increase in the number of sets. This
working platform to the extreme position and imme allowed us to equalize the total amount of training per
diately began to move in the opposite direction. Dur formed by the MC group and the ME group. In addi
ing this training, the amplitude of movements was tion, the duration of the rest periods in this group was
approximately 15% smaller than the amplitude dur reduced from 10 min to 5 min.
ing traditional strength training. Thus, in this group,
Before and after the training cycle, the strength of
the ratio of the times of muscle tension and relax
the trained muscles, as well as the size of the muscles
ation was 100% to 0%.
and muscle fibers of two types, were measured.
In order to evaluate the effect of two types of training Evaluation of hormonal adaptation during the eight
on the size and strength of the muscles involved in the week strength training. The exercises and training proto
movement, before and after the experiment, the 1RM col were the same as in the series with leg extension
for the trained movement was measured. We addi (L series). Leg press 1 RM, the strength of the knee
tionally tested the forcevelocity characteristics of extensor muscles in the isokinetic mode, and the size of
the two main muscle groups involved in this move the m. quadriceps femoris and the two types of muscle
ment, the extensors of the hip and knee joints, in the fibers in its lateral head were evaluated before and after
isokinetic mode, at the angular velocities of 30, 180, the eightweek training.
and 300 deg/s. In addition, we estimated the local per
formance of the knee extensor muscles. The volumes of The effect of strength training of varying intensity on
m. quadriceps femoris and m. gluteus maximus were the biochemical and hormonal blood parameters was
calculated based on magnetic resonance imaging evaluated by the acute response to a traditional training
(MRI); the area occupied by the muscle fibers of the session (comparison of the parameters after a single
first and second types (MFI and MFII) on a cross sec training session with the baseline levels at rest) at the
tion of m. vastus lateralis, as the product of the percent beginning (in the second week) and at the end (seventh
age composition of the fibers of this type in the muscle week) of the training cycle. The lactate concentration,
by the average muscle fiber crosssectional area of this the content of growth hormone (GH), insulinlike
type. growth factor 1 (IGF1), testosterone, and cortisol were
determined in the blood. Before and 18 h after the train
K series. Thirteen young, physically active men ing session, the total activity of creatine kinase (CK)
trained the knee extensors (K series) for 10 weeks. Nine and the activity of its cardiac isoform (MB CK) were
subjects (KC group) used the traditional strength train determined in the blood. The activity of CK from the
ing, and four subjects (KE group) used the lowintensity striated muscle fibers (MM CK) was calculated.
strength training without relaxation. Training protocols
were the same as in the L series. Changes in the expression of genes regulating myogen
esis in response to a single training session. The effects of
The 1 RM of the knee extensor muscles were mea a training session with the knee extension according to
sured before and after 10 weeks of training in the isoki the traditional strength exercise scheme (75% of
netic mode, with the angular velocities of 30, 120, 180, 1 RMtr) was compared to the normal lowintensity
240, and 300 deg/s. exercise (50% of 1 RMtr) and to lowintensity strength
The cross sectional area of the muscle fibers of the exercise without relaxation (50% of 1 RMno relax) that
two types and the area that they occupy were deter were equal in terms of the time of muscle tension and
mined in the same way as in the L series. the range of motion. The study involved young men
During the M series, the experimental group trained adapted to strength training. The changes in gene
with middle intensity. The experiment was organized expression in the muscle sample were determined
according to the following scheme: two groups of ten before, 5 h after, and 22 h after a single training session
subjects each were compared to each other. For the con using the realtime polymerase chain reaction.
trol group (MC group), the subjects of which trained in On the day of the experiment, the subjects had a
the usual way of traditional strength training, each ses standard breakfast; and 1 h after the completion of
sion consisted of seven sets with 5min rest periods training session, they had a standard lunch. Before
between them. During each set, the work with a load of training session and 15 min after the completion of exr
85–90% of 1 RM was performed to exhaustion (6– cise, samples of venous blood were taken to determine
10 reps). In the experimental group (medium intensity the levels of the hormones. During session after the
training without relaxation, ME group) a session con third, fifth, and eighth sets, finger samples of the capil
sisted of five supersets, each of which had four sets. The lary blood were taken (20 μL) to determine the concen
loads were 65–70% of 1 RM; the duration of one set was trations of lactate and glucose. Before the training, as

HUMAN PHYSIOLOGY Vol. 39 No. 5 2013

514 VINOGRADOVA et al.

well as 5 h, and 22 h after the training, using the micro percentage of fibers of this type in the average muscle
biopsy technique [28], a sample of the muscle tissue was fiber crosssectional area of this type.
taken from the m. vastus lateralis. Estimation of hormones and metabolites in the blood
The strength gain and fatiguability of the trained and gene expression in the muscle tissue. The concentra
muscles, the sizes of the muscles and the sizes of muscle tions of lactate, somatotropic hormone, insulinlike
fibers of the two types and the biochemical changes in growth factor 1 (IGFI), testosterone, and cortisol were
the blood were determined before and after the training determined in the blood. Before and 18 h after the
cycle. workout, the total activity of creatine kinase and its car
diac isoenzyme (CK and MB CK, respectively) in the
Evaluation of the forcevelocity characteristics and blood were determined. The activity of CK from the
fatigueability. In order to evaluate the effect of two types striated muscle fibers (MM CK) was calculated as the
of training on the strength gain, before and after the difference between CK and MB CK. The concentra
experiment, one repetition maximum (1 RM) was tions of hormones and enzymes were determined by
determined by the maximum weight that the subject ELISA using DSL (United States) test kits.
could lift once. An additional testing of the forceveloc
ity characteristics of the two main muscle groups that RNA isolation. RNA isolation from the muscle tissue
take part in the movement, the hip and knee extensors, samples was performed using a TRIZOL set (Invitro
was performed using a Pro System 3 dynamometer gen, United States). The complementary DNA was
(Biodex, United States) in isokinetic mode, at the obtained using the Sileks set (Russia).
angular velocities of 30, 120, 180, 240, and 300 deg/s. Polymerase chain reaction. The realtime PCR was
The assessment of local performance (fatigueability) carried out with a RotorGene Q thermocycler (Qiagen,
of knee extensors was carried out in a 45s test with the Germany), using a Sybr Green set (Synthol, Russia). In
maximum isometric force. The dynamics of a decrease the samples, the expression of insulinlike growth factor
in the torque, normalized to 1 RM, was evaluated dur 1 (IGF1Ea) and myostatin genes was determined. The
ing the test. genes of the large ribosomal protein P0 and βactin were
used as reference genes. The changes in the expression
The estimation of the morphometric parameters of the of the target gene were determined according to [30],
muscles and muscle fibers. The volumes of m. quadriceps with the calculation of the efficiency of the reaction for
femoris and m. gluteus maximus were calculated on the the target and reference genes by the standard curve.
basis of MRI. The MRI study was carried out using a
Magnetom 63SP tomograph (Siemens, Germany;
magnetic field, 1.5 T) by the spinecho method in the RESULTS AND DISCUSSION
transverse plane. On each of the images obtained, the
crosssectional areas of m. quadriceps femoris and m. Strength training of the leg extensors (L series). The
gluteus maximus were determined; then, taking into effect of traditional training on the strength of the work
consideration the distances between the shots (17 mm), ing muscles and the size of their muscle fibers (MFs)
the volume of the muscle was calculated. The results was compared to the effect of lowintensity strength
contain the averages of the volume of the muscles of training without muscle relaxation [31]. Strength train
both legs. ing in both modes resulted in an increase in 1 RM of the
leg press. However, in the LC group, where the subjects
The crosssectional areas of the muscle fibers of trained according to the traditional strength training
types I and II were evaluated on the transverse sections protocol, the increase in 1 RM was 34% of the original
of the biopsy samples (m. vastus lateralis), taken using a level; while in the LE group, the increase in 1 RM was
needle biopsy technique [29]. Muscle tissue samples 21% (Table 1). We assume that the greater increase in
were frozen in liquid nitrogen and stored at –70°C. 1 RM was related to the larger training loads. Indeed,
Prior to the analysis, serial sections perpendicular to the the total work performed during the entire training cycle
muscle fibers, with a thickness of 5 μm were made of during traditional strength training was approximately
frozen (–20°C) tissue samples using a microtome twice as much as the total work performed during the
(Leica, Germany). Muscle fibers of types I and II were lowintensity strength training without relaxation
detected by immunohistochemistry using NCLMHCf (Table 1). This difference was due to both the exercise
(a + b) and NCLMHCs monoclonal antibodies intensity and the amplitude of leg extension; as in the
(Novocastra Laboratories). Then, secondary antibodies LE group, each of these parameters was lower than the
labeled with FITS were used. The measurements were respective parameter in the LC group.
made using an Axiovert200 microscope (Carl Zeiss, The maximum strength of the individual muscle
Germany). For each of the tissue samples, the values groups involved in the movement trained were deter
of at least 100 fibers of each type were taken in the mined by isokinetic dynamometry. No differences were
counting. recorded in the gain of the torque of the knee extensor
The area occupied by the muscle fibers of the first muscles; whereas, in the LC group, the gain in the
and second types (MFI and MFII) in the cross section torque of the hip extensor muscles was greater com
of m. vastus lateralis, was evaluated as the product of the pared to that in the LE group in the wide range of angu

HUMAN PHYSIOLOGY Vol. 39 No. 5 2013

 OPTIMIZATION OF TRAINING: NEW DEVELOPMENTS 515

Table 1. The comparison of the effects the traditional highintensive strength training (control groups, C) and training
without muscle relaxation (experimental groups, E) during leg extension series (L series), knee extension series (K series),
and leg extension series, when the group trained with middle intensity (M series)

Group Work, kJ 1 RM gain, % Area of MFI, gain, % Area of MFII, gain, %

L series
LC group (80% 1 RM) 1604 ± 62 34 ± 3.1* 9.4 ± 3.9 23.4 ± 8.3*
LE group (50% 1 RM) 834 ± 44§ 21 ± 2.5*§ 17.7 ± 6.8* 8.2 ± 3.59
K series
KC group (80% 1 RM) 951 ± 60 29 ± 5.8* 6.5 ± 4.3 19.4 ± 5.3*
KE group (50% 1 RM) 400 ± 27§ 24 ± 3.6* 17.7 ± 5.3* 5.6 ± 4.3
M series
MC group (85% 1 RM) 2100 ± 31 33.5 ± 3.9* 5.7 ± 4.5 33.0 ± 5.1*
ME group (65% 1 RM) 2188 ± 33 30.1 ± 4.8* 18.1 ± 6.9* 6.2 ± 3.8
Note: The total work for the entire training cycle; the gains in one repetition maximum (1 RM), and the areas occupied by the muscle fibers
of the first and second types (MFI and MFII) on a cross section of m. vastus lateralis are shown. * Significant gain compared from the
baseline (p ≤ 0.05); § significant difference from this parameter in the control group.

lar velocities (Figs. 1a and 1b). Therefore, we assume i.e., to the middle position between full leg flexion and
that the increase in leg press 1 RM in the LE group was full leg extension in the knee joint.
mainly due to the gain in the strength of the knee exten
sor muscles, rather than to the gain in the strength of the Both types of muscle fibers are involved in the high
hip extensors. intensity muscle contraction [32]. At the same time, it
is well known that the traditional strength training leads
The difference in the gain of the 1 RM of the indi to more severe hypertrophy of type II muscle fibers than
vidual muscle groups is consistent with the MRI data of type I muscle fibers [33–36]. Probably, the reason is
(Fig. 1c). The eightweek training resulted in a compa that highintensity muscular work causes more pro
rable increase (by 10–15%) in the volume of m. quadri nounced metabolic changes in the type II muscle fibers
ceps femoris in both groups. However, an increase in the than in type I fibers. This leads to greater activation of
volume of m. gluteus maximus was recorded only in the the mTORC1 and ERK1/2 signal pathways, which con
LC group; in addition, this increase was greater than the trol protein synthesis [37, 38]. In our study [39], in the
respective increase in the LE group (Fig. 1c). LC group, the area occupied by type II muscle fibers in
Apparently, the differences in the gains in strength the cross section of the working muscle m. vatus lateralis
and volume of the muscles are related to the redistribu was increased by 23%, while for type I fibers, it did not
tion of the load between the muscles during leg exten change, which is consistent with the data in the litera
sion. As noted above, the amplitude of movement in the ture. However, after a lowintensity strength training
groups performing lowintensity strength training was without relaxation, qualitatively different changes in
approximately 15% less than in the group that trained these parameters were found: the area occupied by type
according to the traditional scheme. It is well known II muscle fibers did not change, while the area occupied
that the strength of the skeletal muscles depends on by type I muscle fibers increased by 18% (Table 1). We
their length (the strength–length relationship). For the hypothesized that the marked metabolic stress in the
hip extensors, the optimal muscle length (the length at muscle fibers could be one of the factors that initiate the
which a given muscle develops the maximum force) is process of hypertrophy. It is difficult to achieve the
recorded in the position of a flexed hip [31]. In the LE marked metabolic shifts in type I muscle fibers [40],
group, the extensor muscles of the hip did not achieve since the fibers of this type have a high oxidation poten
their optimal length during exercising due to the lower tial. However, it was shown that, during isometric con
amplitudes of their movement. As a result, during low tractions (under the conditions of the restricted blood
intensity strength training without relaxation, in the L flow caused by muscle tension) with intensities of 25,
series, the hip extensors developed smaller strength, 50, and 75% of 1 RM, the maximal lactate concentra
and, consequently, they obtained a smaller training tion in m. vastus lateralis at the exhaustion was recorded
effect than during the traditional training. For the knee when the intensity was 50% of 1 RM, with the largest
extensors, the decrease in the amplitude did not affect concentration of lactate found in type I muscle fibers
the biomechanics of contraction, as the optimal length [41, 42]. Apparently, this is due to the fact that, accord
of this muscle group corresponds to 110 degrees [31], ing to the principle of dimension, work of this character

HUMAN PHYSIOLOGY Vol. 39 No. 5 2013

516 VINOGRADOVA et al.

40 relaxation, with an intensity of 50% of 1 RM, as well as

(a) * during isometric exercises, primarily, type I muscle
fibers are involved in the performance, which allows the
Torque, gain, %
30 *
* § marked metabolic shifts in them to be achieved at
§ § exhaustion. Note that, in the LE group, the termination
20
of each set was accompanied by severe subjective fatigue
of the working muscles. At the same time, the concen
10 tration of blood lactate, which is the marker of the acti
vation of glycolysis, during the training session in the
0 LE group was higher than that in the LC group (see
below).
(b) 40
The increase in the resistance of the trained muscles
LC group to fatigue in the 45s maximal isometric test compared
Torque, gain, %

30 LE group to the traditional strength training is one of the expected

effects of the lowintensity strength training without
20 relaxation. In assessing the resistance to fatigue in the
* * * static test, it was found that, in the LC group, there was
10 * a decrease in this parameter after the eightweek train
ing cycle, while in the LE group, it did not change com
0 pared to the baseline levels before the training [39].
30 180 300
Angular velocity (deg/s) Thus, it has been shown that there was an increase in
the muscle mass and strength after eight weeks of low
(c) 30 intensity (50% of 1 RM) strength training without
Muscle volume, gain, %

relaxation. Note that the increase in the muscle mass

25
*
after eight weeks of lowintensity strength training with
20 * § out relaxation is related to the hypertrophy firstly of type
15
I muscle fibers, while the traditional strength training
* leads to the hypertrophy preferentially of type II muscle
10 fibers. In the this study, the effects of exercises in the L
5 and K series were compared. Note that studying leg
extension is associated with a number of methodologi
0 cal restrictions for analysis, which is due to the specifics
m. quadriceps femoris m. gluteus maximus of biomechanics and the control over complex motor
40 *
actions, with the redistribution of efforts between differ
(d) KC group ent muscle groups, etc. With this kind of training, the
35 KE group
Torque, gain, %

30
* objective assessment of a variety of physiological and
* * morphological responses at the level of individual mus
25 *
cle groups may be difficult. Therefore, the data obtained
20 during a 10week training of a knee extensors are of par
15 * * * ticular interest.
10 Strength training of the knee extensors; K series. The
5 effects of the traditional strength training with an inten
0 sity of 80–85% of 1 RM (control group, KC group)
30 120 180 240 300 were compared to the effects of lowintensity strength
Angular velocity (deg/s) training without relaxation, with an intensity of 50% of
1 RM (experimental group, KE group). The total work
Fig. 1. The changes in the torque at different angular veloc
ities of the hip (a) and knee (b) extensor muscles; (c) the done during a 10week training cycle differed in the KC
crosssectional areas of m. quadriceps femoris and m. glu group and KE group by more than two fold (Table 1). At
teus maximus estimated by MRI in the L series; and (d) the the same time, the gains in 1 RM of the knee extensor
changes in the torque at different angular velocities in the K muscles in the trained exercise did not differ, being 29
series. * Significant (р ≤ 0.05) gains compared to the base and 24% in the KC group and KE group, respectively
line levels; § significant (р ≤ 0.05) difference of experimental
(E) group from the respective parameter in the control (C) (Table 1).
group. For the abbreviations here and in Fig. 2, see the text. Analysis of the gains in 1 RM during training in the
L series and K series showed significant differences. In
should primarily engage type I muscle fibers [32], in the K series, the gain in 1 RM after traditional strength
which, due to the limited blood flow, the activation of training is greater than that after lowintensity training
glycolysis occurs. This provides grounds to assume that, without relaxation (35% vs. 21%); while in the K series,
during the lowintensity strength training without no differences between the groups were recorded (29%

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 OPTIMIZATION OF TRAINING: NEW DEVELOPMENTS 517

vs. 24%). Apparently, these differences are related to the The total work performed during the 8week training
features described above of the biomechanics of the period was 2100 ± 31 kJ for the MC group and 2188 ±
muscles during leg and knee extension, namely, the 33 kJ for the ME group, so it did not differ between the
redistribution of the load between the muscles. groups (Table 1). Gains in the leg press 1 RM over
8 weeks in the MC group and ME groups were 34% and
Assessing the effect of a 10week training of the knee 30%, respectively, i.e., no differences between the
extensors on the area occupied by muscle fibers of dif groups were recorded (Table 1).
ferent types in the cross section of m. vastus lateralis
showed similar gains in type II muscle fibers in the KC Assessing the physiological and morphological
group and type I muscle fibers in the KE group parameters of the knee extensor muscles showed the
(Table 1), which is in good agreement with the results same gains in 1 RM in isometric mode (8.0 ± 3.2% in
obtained during training in L series. the ME group and 7.2 ± 3.5% in the MC group) and the
same gains in the volume of m. quadriceps femoris eval
Thus, based on the comparison of the effects of the uated by MRI (9.8 ± 1.6% in the ME group and 6.8 ±
prolonged training of the knee and leg extensor mus 2.4% in the MC group).
cles, we conclude that (1) the degree of hypertrophy of
the knee and hip extensors during training in leg exten As in the group that trained with low intensity with
sors depends on the biomechanics of the exercise; (2) as out relaxation, in the group that trained with middle
a result of the redistribution of the load between the intensity without relaxation, an increase (18.1 ± 6.9%)
muscles, the strength gain of the knee extensor muscles was found in the area occupied by type I muscle fibers
during leg extension lowintensity training without in the cross section of m. vastus lateralis, and no
relaxation is lower than that during traditional strength increase was found in the area occupied by type II mus
training; (3) in the case of training of knee extensors, cle fibers (Table 1). In contrast, in the group that used
lowintensity strength training without relaxation can traditional strength training, there was an increase in
result in the same increase in strength of knee extensors the area occupied by type II muscle fibers (by 33.0 ±
muscle as the traditional strength training; (4) low 5.1%), and no changes were found in the area occupied
intensity training without relaxation of the knee exten by type I muscle fibers. Thus, training without relax
sor muscles, as well as training of the leg extensors, ation with an intensity of about 65–70% of 1 RM can
results in an increase in muscle strength, mainly due to increase the strength to the same extent as traditional
the hypertrophy of type I muscle fibers. highintensity strength training.
When comparing the effects of strength training and
So far, we compared the effects of traditional lowintensity training without relaxation, the question
strength training (80–85% of 1 RM) and lowintensity about the mechanisms of the gain in the muscle mass
training (50% of 1 RM) without relaxation. In the LE naturally arises.
group, the increase in the strength was smaller than in
the group that trained with a high intensity, which could An increase in the muscle mass and strength during
be due to lower intensity and volume of the work per lowintensity training without relaxation is related to hor
formed. We assume that, during training with incom monal adaptation. As noted above, muscle hypertrophy
plete relaxation, a slight increase in the intensity (from can be activated by mechanical and metabolic factors.
50% of 1 RM to 65–70% of 1 RM) and an increase in Both kinds of factors can activate protein synthesis
the work performed during a training session (to the lev either directly, by affecting the muscle fiber, or indi
els of traditional strength training) could provide an rectly, through increased secretion of anabolic hor
increase in strength comparable to their increase during mones. We attempted to answer the question on which
traditional highintensity strength training. of these factors (mechanical or metabolic) is more
important for triggering the hormonal response to mus
Strength training of mediumintensity without relax cle activity, and determine the mechanisms of hyper
ation (M series). The scheme of the experiment com trophic changes in the muscle tissue when using differ
paring two groups of subjects was described above. The ent types of training.
subjects trained leg extension. In the control group
(MC group), which practiced traditional strength train There is evidence that an increase in the concentra
ing, the load was 85–90% of 1 RM. In the experimental tion of the growth hormone is related to the activity of
group (medium intensity training without relaxation, Ia muscle afferents [43]. However, many authors
ME group), the load was 65–70% of 1 RM. The work attribute the increased concentrations of anabolic hor
out of the ME group was increased compared to the mones to metabolic changes in the working muscles [8,
series previously described due to the greater intensity 9, 44, 45]. Indeed, it is shown that limb ischemia poten
of the load (65–70% of 1 RM versus 50% of 1 RM in the tiates the hormonal response when performing strength
preceding experiment) and the increase in the number [20] and aerobic exercise [10, 20].
of sets. This allowed us to equalize the total work per In our study [46], we hypothesized that the restric
formed by the MC group and ME group. Furthermore, tion of the blood supply of the working muscles during
in the ME group, the duration of the rest periods was lowintensity training without relaxation serves as a
reduced from 10 min to 5 min. stimulus for the secretion of anabolic hormones. The

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518 VINOGRADOVA et al.

Table 2. The changes in the blood concentrations of lactate, somatotropic hormone (GH), insulinlike growth factor
(IGF1), testosterone, cortisol, and the activity of muscle creatine kinase (MM CK) in response to a single training session in
week 2 and week 7 of traditional highintensive strength training (80% of 1 RM, C group) and lowintensity (50% of 1 RM)
training without muscle relaxation (E group)

C group E group

Parameters week 2 week 7 week 2 week 7

before after before after before after before after

Lactate, mM 2.2 ± 0.2 8.5 ± 0.6* 1.8 ± 0.2 12.3 ± 1.0* 2.4 ± 0.3 13.2 ± 1.8* 2.5 ± 0.3 14.5 ± 1.4*
MM CK, 176 ± 30 435 ± 76 106 ± 24 255 ± 42 145 ± 32 283 ± 61 123 ± 9 182 ± 11
MU/L
GH, ng/mL 0.22 ± 0.02 3.47 ± 0.76* 0.20 ± 0.05 6.30 ± 2.82* 0.27 ± 0.03 9.09 ± 1.37* 0.20 ± 0.03 8.29 ± 1.41*
IGF1, 277 ± 39 259 ± 29 276 ± 27 266 ± 30 320 ± 50 350 ± 47* 304 ± 17 372 ± 30*
ng/mL
Testosterone, 13.89 ± 2.43 14.24 ± 2.78 12.15 ± 1.74 13.89 ± 2.08* 9.72 ± 1.04 11.81 ± 2.08 7.99 ± 1.74 8.68 ± 2.08
nmol/L
Cortisol, 195 ± 19 259 ± 37 253 ± 22 341 ± 53 154 ± 18 264 ± 29* 187 ± 14 358 ± 30*
nmol/L
* Significant (p ≤ 0.05) gains compared to the baseline levels.

training protocols were the same as in the L training in training cycle was 73% greater in group C (traditional
the study described above [39]. strength training) than that in group E (lowintensity
It was found that the traditional strength training training without relaxation) (258 ± 69 IU/L vs. 149 ±
(85% of 1 RM) and lowintensity training without 36 IU/L), which is likely to be due to the greater train
relaxation (50% of 1 RM) cause different changes in the ing load in group C (85% of 1 RM vs. 50% of 1 RM,
blood lactate concentration and in the activity of the respectively). Probably, in the subjects of group E, the
muscle isoform of creatine kinase (MM CK). Lactate is restriction of blood flow in the working muscle did not
the final product of glycolysis. Since, during strength potentiate the damage of the membranes of muscle
exercise, the energy supply is provided mainly by the fibers. This is consistent with the finding that occlusion
anaerobic pathway and the level of blood lactate indi of the working limb does not increase the activity of CK
rectly shows the intensity of metabolic processes in the in the blood: one day after lowintensity strength train
working muscles. As expected, the restriction of the ing (20% of 1 RM) with the occlusion of working mus
blood supply of the working muscles was accompanied cles (214 mm Hg), the level of CK was the same as in the
by a marked increase in the concentration of the lactate control group exercising without occlusion [20].
in the blood. At the beginning of the eightweek training Comparing the changes in the concentrations of lac
cycle (during week 2), after the training session, the tate and MM CK during two types of training load used
blood lactate concentration during traditional strength in our study [46], we suggest that the traditional strength
training (C group) increased by 6.3 mmol/L; however, training causes greater mechanical damage to the mus
after lowintensity strength training without relaxation cle fibers, increasing the release of MM CK in the
(E group), it increased by 10.8 mmol/L (Table 2). In the blood. At the same time, after the training session with
final period of the training cycle, the differences incomplete relaxation, intensified glycolysis leads to a
between the groups became less marked. Thus, the con significant accumulation of lactate in the blood. Thus,
centration of blood lactate after a session of training traditional training generates predominantly
with incomplete relaxation, despite the lower volume of mechanicrelated stimuli that can trigger anabolic pro
work, was greater rather than less than that after the ses cesses in muscle fibers, whereas after training with
sion of training in the traditional way. incomplete relaxation, metabolic changes develop
It is well known that the release of musclular CK in more intensely. If we assume that the main feature of
blood indirectly reflects the extent of damage of the cell exercise with incomplete relaxation is the restriction of
membrane of working muscles and depends on the the blood supply in working muscles, then we should
load. For example, during the eccentric exercise, the consider the data on training under the conditions of
increase in activity of CK is greater than that during the occlusion in our discussion on the mechanisms of
concentric exercise [47–49]. In our study [46], an hypertrophic response. For example, according to [21,
increase in the activity of MM CK during week 2 of the 22], a fourweek aerobic exercise (cycling 4 times a

HUMAN PHYSIOLOGY Vol. 39 No. 5 2013

 OPTIMIZATION OF TRAINING: NEW DEVELOPMENTS 519

week for 45 min, load ~20% of 1 RM) in a hermetic cates the high physiological costs of these types of train
chamber with extra pressure on the lower half of the ing (Table 2).
body of 50 mm Hg leads to an increase in the crosssec Thus, lowintensity (50% of 1 RM) exercise without
tional area of the trained muscles, and an increase in the relaxation leads to higher levels of secretion of GH and
crosssectional area of the muscle fibers predominantly IGF1 in the blood than exercises with larger external
of types I and IIB. The authors attribute hypertrophy to loads (85% of 1 RM) performed in the traditional way.
both an increase in contractile proteins, and an increase Probably, this phenomenon is caused by more intensive
in glycogen and hyperhydration. Similar results were accumulation of metabolites in the muscle during train
obtained in the study on elite rugby players performing ing with incomplete relaxation due to the reduced blood
an eightweek ischemic (cuff pressure, 200 mm Hg) flow, which is inevitable with this type of training, as
lowintensity strength training (twice a week, 4 sets, well as under the conditions of muscle ischemia. Note,
50% of 1 RM) [50]. The gains in the dynamic force, and however, that the mechanisms of metabolic reflectory
the cross section (MRI) of the knee extensors in this posteffect stimulation of the secretion of anabolic hor
study were 14 and 15%, respectively. Comparing the mones require further research. In studies on the hor
data on the changes in the specific force and elec monal response to strength training, the results are
tromyographic activity, the authors come to the conclu mixed. West et al. studied the effects of different levels of
sion that the increase in the muscle volume is mainly endogenous hormones in the blood on the strength gain
due to contractile proteins. and muscle hypertrophy [53]. The authors concluded
that an increase in the levels of the growth hormone and
The increase in the secretion of GH and insulinlike IGF1 in the blood caused by the strength exercise is
growth factor 1 (IGF1) can be considered among the not the main stimulus for the activation of a hyper
factors responsible for the hypertrophic effect due to trophic process in the working muscles. The regulation
training against the restriction of blood supply. In our of the synthesis of muscular protein, in their view,
study [46], the concentration of GH in the blood after a depends mainly on the intracellular mechanical and
training session with incomplete relaxation was higher; metabolic stimuli.
although the mechanical effect on the muscle fibers was
much smaller than during the traditional strength train Effects of highintensity strength training and low
ing (Table 2). It is important that training without relax intensity strength training without relaxation of muscles on
ation provided a high GH response throughout the the expression of genes regulating myogenesis. In the past
entire training cycle. A significant increase in the blood decade, molecular biological techniques have been
GH was shown previously during aerobic and strength widely used to study the mechanisms underlying muscle
exercise with venous occlusion [10, 20]. According to adaptation to various physical activities and to deter
the authors, this is due to the activation of afferents III mine the efficiency of different training programs.
and IV in response to the local accumulation of metab Analysis of gene expression in muscle cells enables us to
olites in the muscles working with limited blood supply evaluate, individually for each subject, the efficiencies
(metaboreflex mechanism). of different training protocols based on the results of a
single training session. It was shown that a single session
The main system mediator of the anabolic effects of of highintensity strength training alters the expression
the growth hormone is the IGF1. It is well known that of myogenic regulatory genes, such as IGF1, MGF,
it is secreted systemically by the liver cells stimulated by MyoD, Myogenin, and Myostatin, during the day of
the growth hormone and autocrinically, by the skeletal recovery and longer [54, 55]. It was found that low
muscle fibers during their intense contractile activity intensity training (20% of 1 RM) with limited blood
[51]. Thus, an increase in IGF1 in the blood was flow leads to changes in the expression of the genes
recorded after highintensity strength exercise [12, 14]. studied; whereas in the control group training of the
In our study [46], increases in the levels of IGFI in the same intensity, the changes in the expression may be
blood (during week 2, as well as week 7) were recorded absent [56, 57]. This is due to the fact that, during tradi
only after the training session with incomplete relax tional strength exercise, principally greater external
ation. After a traditional strength exrcise (characterized loads are used (70–90% of 1 RM); whereas, in the cited
by much greater intensity of mechanical loads) no studies, the load of 20% of 1 RM was used in control
changes in the concentrations of IGF1 were detected group, which may be too small to cause the changes in
(Table 2). Note that an increase in IGF1 after exercise the expression of myogenic regulatory genes in the mus
with incomplete relaxation occurred against the back cles of physically active people. As mentioned above, in
ground of an increase in the secretion of GH (Table 2). order to achieve marked metabolic stress in the working
muscle during exercise with restricted blood flow, the
Lowintensity exercise with restricted blood flow load should be at least 30–40% of 1 RM [25]. There
caused no increase in the content of testosterone in the fore, we attempted to compare the changes in the
blood (Table 2), which is consistent with the data of expression of myogenic regulatory genes after strength
other authors [52]. At the same time, we note that low exercise performed without muscle relaxation (50%
intensity exercise without relaxation resulted in 1 RMno relax), to the changes after the traditional high
increased cortisol secretion, which, apparently, indi intensity strength exercise.

HUMAN PHYSIOLOGY Vol. 39 No. 5 2013

520 VINOGRADOVA et al.

(a) According to the current view, in adult humans, the

3 IGF1 protein secreted by the skeletal muscles may
75% 1 RMtr affect muscle hypertrophy, mainly, due to the regulation
50% 1 RMno relax of the differentiation and fusion of satellite cells with
50% 1 RMtr existing muscle fibers [59, 60]. In addition, the suppres
IGF1 Ea mRNA

2 sor effect of IGF1 on the degradation of muscle pro

teins through signaling pathway AktFOXO [61] and its
possible role in the regulation of the rate of protein syn
thesis through activation of AktmTORC1 is discussed
1 [60, 62]. In our study the expression of the muscle iso
form of the insulin growth factor (IGF1Ea) in the
recovery period did not change (Fig. 2).
The training regimes compared influenced the
expression of the Myostatin gene, which is a negative
0 10 20 regulator of the muscle growth. Five hours after exercise
(b) without relaxation, the content of myostatin mRNA
decreased by fourfold; in addition, it remained at that
1 level until 22 h after exercise (Fig. 2). After highinten
#
sity strength exercise, the content of myostatin mRNA
Myostatin mRNA

continuously decreased for 22 h of the recovery (in

total, by 20fold). Note that, after the lowintensity tra
* ditional exercise, this expression of the gene did not
0.1 change. The decrease in the expression of myostatin is
related to the increased activity of signaling kinases of
** AktmTORC1 pathway [63]; on the other hand, the
decrease in the expression of myostatin can decrease
activity of the FOXO transcription factors, which stim
0 10 20 ulate the expression of E3 ligases mRNA, and inhibit
Time, h the differentiation of satellite cells [64, 65].
Thus, it has been shown that different modes of
Fig. 2. The expression of mRNA of IGF1Ea (a) and myo strength exercise result in different changes in the
statin (b) normalized to the baseline levels, after a single expression of myogenic regulatory genes: highintensity
strength training: traditional highintensity strength training
(75% of 1 RMtr), lowintensity training without relaxation strength exercise decrease of the myostatin expression
(50% of 1 RMno relax), and traditional lowintensity strength higher than lowintensity training without relaxation.
training (50% of 1 RMtr). 0 h, the time of the termination of Obviously, in order to characterize in detail the effects
a training session. * Significant changes (р ≤ 0.05) compared of different types of strength exercise, we should assess
to the baseline levels; # significant difference (р ≤ 0.05) from the changes in the rate of myofibrillar protein synthesis
the respective parameter of 75% of 1 RM and 50% of 1 RM
and activation of signaling pathways that regulate the
rate of protein synthesis. These tasks should be solved in
To eliminate the effect of redistribution of the load future studies.
between different muscle groups during the exercise and
the impact of the increased secretion of anabolic hor CONCLUSIONS
mones on intracellular signaling, the training of knee
extensors was chosen. It is well known that strength Comparison of the efficiency of different types of
exercises involving low muscle mass do not lead to strength training showed that, after low intensity (50%
increased secretion of anabolic hormones [53, 58]. In of 1 RM) strength training without relaxation of the
working muscles, the gains in strength capacities and
the present study, during the training, the concentration
the size of the trained muscles were not as large as those
of lactate in the capillary blood increased slightly (to after traditional highintensity (80–85% of 1 RM)
3.1 ± 0.5 mmol/L) during lowintensity training (50% strength training. Muscular hypertrophy caused by the
of 1 RMtr); and it increased to 6.3 ± 0.6 mmol/L after traditional strength training and by training without
a highintensity training session. After lowintensity relaxation is, apparently, achieved in different ways.
training without relaxation, the concentration of lactate During the traditional strength training, mechanically
reached the highest levels (7.4 ± 0.5 mmol/L). As related stimuli, which trigger anabolic processes within
expected, the content of anabolic hormones: testoster the muscle fibers, are predominantly generated. During
one and IGF1 did not change after the training ses training without relaxation, there is more pronounced
sions in either of the exercises. The content of GH activation of anaerobic reactions in the muscle, which
increased slightly only after highintensity strength by the metaboreflex leads to a more pronounced
exercise. increase in the secretion of GH, IGF1, and cortisol in

HUMAN PHYSIOLOGY Vol. 39 No. 5 2013

 OPTIMIZATION OF TRAINING: NEW DEVELOPMENTS 521

the blood. Highintensity strength exercise and low 8. Lin, H.S., Wang, W., Wang, R.Y., et al., Stimulatory
intensity strength exercise without muscle relaxation effect of lactate on testosterone production by rat leydig
are characterized by different dynamics of the expres cells, J. Cell Biochem., 2001, vol. 83, p. 147.
sion of the myogenic regulatory gene Myostatin. The 9. Lu, S.S., Lau, C.P., Tung, Y.F., et al., Lactate and the
differences in the triggering mechanisms are combined effects of exercise on testosterone secretion: evidence for
with the differences in the features of hypertrophic the involvement of a campmediated mechanism, Med.
changes under the influence of two types of training. Sci. Sports Exerc., 1997, vol. 29, no. 8, p. 1048.
During traditional strength training, there is a preferen 10. Viru, M., Jansson, E., Viru, A., et al., Effect of restricted
tial increase in the crosssectional area occupied by type blood flow on exerciseinduced hormone changes in
II muscle fibers; during training without relaxation, healthy men, Eur. J. Appl. Physiol. Occup. Physiol., 1998,
there is a preferential increase in the crosssectional vol. 77, no. 6, p. 517.
area occupied by type I muscle fibers. 11. Ahtiainen, J.P., Pakarinen, A., Alen, M., et al., Muscle
hypertrophy, hormonal adaptations and strength devel
This study can be useful in the practice of sports, opment during strength training in strengthtrained and
especially those with high requirements for the strength untrained men, Eur. J. Appl. Physiol., 2003, vol. 89,
and endurance of the working muscle. In addition, no. 6, p. 555.
training with relatively low loads preserves the ligamen 12. Kraemer, W.J., Marchitelli, L., Gordon, S.E., et al.,
tous from injuries. This feature allows this training Hormonal and growth factor responses to heavy resis
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ACKNOWLEDGMENTS Physiol., 1998, vol. 78, no. 1, p. 69.
This study was supported by the Basic Research Pro 14. Rojas, V.S., Knicker, A., Hollmann, W., et al., Effect of
gram of the Russian Academy of Sciences (project resistance exercise on serum levels of growth factors in
humans, Horm. MeTable Res., 2010, vol. 42, no. 13,
no. 6013/1) and the Russian Foundation for Basic p. 982.
Research (project no. 120401668a). The study was
carried out with the use of equipment purchased using 15. Fry, C.S., Glynn, E.L., Drummond, M.J., et al., Blood
flow restriction exercise stimulates mTORC1 signaling
the funds of the Development Program of Moscow and muscle protein synthesis in older men, J. Appl. Phys
State University. iol., 2010, vol. 108, no. 5, p. 1199.
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