APPLIED SCIENCES: PHYSICAL FITNESS AND PERFORMANCE

Muscular adaptation and strength during

the early phase of eccentric training:
influence of the training frequency
SORICHTER, STEPHAN; MAIR, JOHANNES; KOLLER, ARNOLD; SECNIK, PETER;
PARRAK, VOJTECH; HAID, CHRISTIAN; MÜLLER, ERICH; PUSCHENDORF, BERND
Author Information
Medicine & Science in Sports & Exercise: December 1997 - Volume 29 - Issue 12 - p 1646-
1652

Abstract
Muscular adaptation and strength during the early phase of eccentric training: influence of the
training frequency. Med. Sci. Sports Exerc., Vol. 29, No. 12, pp. 1646-1652, 1997. We
investigated the effects of different training frequencies on maximum isometric voluntary
contraction(MVC) force and plasma concentrations of muscle proteins during the early phase
of eccentric training. MVC and plasma concentrations of creatine kinase(CK) and slow-twitch
skeletal (cardiac beta-type) myosin heavy chain (MHC) fragments were measured before and
4 and 7 d after performing the first and last training task. Training tasks, which comprised 70
high-force eccentric contractions involving the thigh muscles (single leg), were performed
under supervision in three groups (A, B, C) at the beginning and at the end of the study period
(7 wk). In addition, groups A (N = 10) and B(N = 10) trained during the study period starting
1 wk after the first training task. Group A performed one training task once a week for 5 wk
and group B (N = 10) twice a week for 2 wk and three times a week during the subsequent 3
wk. In all three groups the first training task resulted in delayed CK and MHC peaks and
decrements in MVC, which were comparable (P > 0.05). Only training regimen B resulted in
a significant increase in the MVC. Compared with the first training task training regimens, A
and B significantly (P < 0.01) reduced the increase in serum muscle protein and muscle
function impairment. The responses to the last training task did not differ significantly
between groups A and B. In group C the responses after the second training task did not differ
significantly from those observed after the first task. Our results suggest that, compared with
group A, additional eccentric exercise in group B is the essential basis for the increase in
muscle strength during the early phase of eccentric training without further benefits for
muscular adaptation. In group C we found no muscular adaptation.

Training protocols involving maximal eccentric contractions are established methods for
improvement of muscle strength. Training of the muscle strength is based on two different
adaptation processes: muscle fiber hypertrophy and neurological adaptation(3,12,13,29).
Additionally, changes in the quality of protein without hypertrophy is one of the primary
adaptation processes during the early phase of heavy-resistance muscle strength training(32).

Few studies exist on the question of the optimal training frequency(24) in the early phase of
eccentric heavy-resistance training. Until now, only the efficiencies of specially designed
training protocols were investigated(12,15,16,32). Most of these training protocols included a time
period of 6-8 wk with 2-3 training tasks per week (15,16,32). Earlier investigations on force
training showed that pure eccentric or a combination of eccentric and concentric exercise
generate a high training stimulus which improves the rapid force development (16) as well as
the individual strength (11,30) and was mostly quantified in previous studies as the maximal
isometric voluntary contraction(MVC) force (35). It is well established that strenuous
unaccustomed eccentric exercise may result in increases in muscle protein serum levels (i.e.,
creatine kinase (CK), myosin heavy chain fragments (MHC), muscle soreness and stiffness,
prolonged strength loss, and morphological changes (7,22,26) reflecting exercise-induced muscle
damage. This eccentric exercise-induced skeletal muscle damage induces repair or
regeneration processes(31) that result in a temporary muscular adaptation. On the other hand,
during this time period the ability to generate muscle strength is reduced and prevents a
successful training of technical abilities in performing sports-specific movements. Therefore,
it is necessary to know the optimal frequency of training tasks in the early phase of eccentric
strength training, which leads to an increase in MVC without extended regeneration periods.

The purpose of the present study was to investigate the effects of three different training
frequencies during the early phase of eccentric strength training. We chose an eccentric
exercise protocol involving the thigh muscles that is known to produce muscle damage at the
beginning, and then subjects trained with it to evaluate its efficiency and muscular adaptation.
The effects of the different training regimens were assessed by the improvement of MVC.
Similar to earlier studies(4,7,22,23), the adaptation of the trained muscles was monitored by
measuring plasma concentrations of muscle proteins and assessing muscle soreness.

MATERIALS AND METHODS

Subjects. Thirty healthy male volunteers (physical education teacher trainees) ranging in age
from 21 to 25 yr were recruited from the Department of Sports Sciences (University of
Innsbruck). All subjects had no physical limitations to exercise and were not involved in any
unilateral leg training. The risks and benefits of the study were explained and written
informed consent was obtained from each participant. The study was approved by the
institutional review board of the local institution. Subjects were randomly assigned to one of
three groups (see Table 1). All subjects were instructed to refrain from unaccustomed exercise
during the course of the study starting 48 h before the first exercise session.

Training design. Exercise was conducted in a sitting body posture on an exercise rack
specially designed to elicit the required eccentric action of the musculus quadriceps
femoris (22,31). All subjects were tested for their maximal strength generation of the investigated
leg with the knee held at an angle of 1.75 rad (3.14 rad correspond to full extension of the
knee). Subjects then had to hold their knees at an angle of 2.62 rad, when a special trigger
mechanism suddenly released 150% of the maximal voluntarily generated force. Subjects
were instructed to straighten their knee against the pressure of this weight. But given the
arrangement they could not help bending their knee, although they tried to resist. A pulley
system allowed the researcher to bring the weight to the starting position without any loading
concentric exercise of the investigated leg. After warming up each subject performed a single
bout of eccentric exercise using only the nondominant leg. The training task consisted of
seven sets of 10 eccentric contractions of the quadriceps femoris muscle group. Each
contraction lasted 1-2 s, with 15 s of rest between contractions. The seven sets were each
separated by 3 min of rest. To evaluate the training effects during the early phase of eccentric
training, a training period of 6 wk was investigated. Under supervision and with adjustment to
the increases in muscle strength, three groups (A, B, C; N = 30) performed the first training
task at the beginning and the last training task after 7 wk at the end of the study period.
Groups A (N = 10) and B (N = 10) additionally trained during the study period starting 1 wk
after the first training task. Group A trained once a week for 5 wk and Group B (N = 10) twice
a week for 2 wk and three times a week during the subsequent 3 wk.

Warm-up. The 10-min warm-up consisted of 5 min running because of the individual aerobic
threshold, followed by 5 min of stretching the leg muscles, and finishing up with three series
of eight knee bends.
(7)
Muscle soreness evaluation. Similar to earlier studies , muscle soreness and pain in all
subjects were assessed by questionnaire before, and 1, 2, 3, 4, and 5 d after the first and last
training task. On a scale of 0 (normal) to 10 (very sore) subjects rated the perceived soreness
of the loaded quadriceps muscle.

Isometric muscle function test. The ability to generate MVC of the knee extensors was
monitored before doing the first and the last eccentric exercise bout and 4 and 7 d after
finishing, respectively. The procedure for investigating muscle function was adapted from a
method described by Edwards et al. (8). After warm-up the subject was seated in an adjustable
straight-backed chair, and a not extensible taut strap was passed around the ankle just above
the level of the lateral malleolus. A seat belt was passed around the waist to firmly secure the
pelvis. An 8-cm steel chain connected the strap with a load cell under the chair so that the
knee angle was 1.57 rad. The load cell (Type V 2a, Hottinger-Baldwin, AT) was connected to
a 2A/2D amplifier/transducer (DMC plus, Hottinger-Baldwin, AT) and the forces were
recorded and displayed on the analog-to-digital data acquisition software package. Each
subject performed three isometric MVCs of both quadriceps femoris, with a break of 1 min
between each. The highest value achieved in the repetitions was used as the criterion score.
The test-retest reliability was 0.92.

Blood analysis. Blood samples were drawn from a superficial vein of the forearm and
collected in ethylenediaminetetraacetate (EDTA) coated tubes (Sarstedt, Nümbrecht,
Germany) and immediately centrifuged. Samples were withdrawn immediately before
performing the first and last eccentric training task and 4 and 7 d after finishing, respectively.
Plasma CK activity was measured on the same day. Aliquots of plasma samples for MHC and
cardiac troponin I measurements were subsequently frozen and stored at -20 °C until assayed.
All measurements were done in duplicate.

CK activity. CK (molecular weight 88000) is a key enzyme of muscular metabolism that

exists predominantly as a soluble sarcoplasmic protein in muscle fibers. CK is found in all
types of skeletal muscle fibers in similar concentrations. Its half-live time in the general
circulation is approximately 15.5 h (18). CK activities were measured at a temperature of 25 °
Celsius by means of a N-acetylcysteine activated, optimized UV test obtained from Merck
(Darmstadt, Germany). For men the upper limit of the reference interval of CK is 80 U·L-1.
The intra- and interassay coefficients of variation (CV) were 8.2% and 10.6%, respectively.

Myosin heavy chain. Myosin is a hexameric structurally bound contractile protein containing
4 light and 2 heavy chains (molecular weight 230000). MHC can be cleaved into its
subfragments by enzymes. The rod portion can be further degraded to form light meromyosin
and subfragment 2 (molecular weight 51000; 33). Concentrations of MHC fragments were
measured by an immunoradiometric assay (E.R.I.A. Diagnostics Pasteur, Marnes la Coquette,
France). This sandwich assay uses a pair of monoclonal antibodies primarily raised against
two different epitopes on subfragment 2 in the rod of human ventricular beta-type heavy
meromyosin. The antibodies used have been described in detail elsewhere (20,21). Briefly, owing
to the strong structural similarity of beta-type cardiac MHC and MHC of slow-twitch skeletal
muscle fibers (5,36), both antibodies react strongly with human slow skeletal MHC. The
affinities of the antibodies to slow skeletal muscle myosin were identical to those to beta-type
cardiac MHC (20). By contrast, the antibodies do not significantly react with cardiac alpha-type
MHC, nor with MHC of human fast-twitch skeletal muscle fibers or of any human smooth
muscle. Thus, the assay recognizes very well MHC fragments of human beta-type and slow
muscle MHC. The monoclonal antibodies of the assay recognize MHC subfragment 2 in the
whole molecule and as proteolyzed fragments. The upper limit of the reference interval of
MHC in plasma is 300 μU·L-1. The detection limit of the assay is 10 μU·L-1. The intra- and
interassay CVs were 3.8% and 7.2%, respectively. One μU/L corresponds to 1μg·L-1(20).

Cardiac troponin I assay. Troponin I (TnI) is a contractile protein of thin filaments in

striated muscle fibers. It is part of the troponin-tropomyosin complex and exists in three
different isoforms, one for slow-twitch skeletal muscle, one for fast-twitch skeletal muscle,
and one for cardiac muscle fibers (34). A highly specific immunoenzymometric assay (E.R.I.A.
Diagnostics Pasteur) developed by Larue et al. (19)was used to detect circulating cardiac
TnI(molecular weight 24000 Da) in plasma. This assay does not cross-react with TnI of
human skeletal muscle. No cardiac TnI (<0.1 μg·L-1) could be detected in the plasma of
healthy volunteers or blood donors.

Statistics. The BMDP statistical software package was used for data analysis. The influence
of training on responses to exercise were tested by ANOVA and ANCOVA with repeated
measures and Greenhouse-Geisser adjustment of the P-value. Differences between groups
were assessed by one-way ANOVA with a Newman-Keuls post hocanalysis. The paired
Studentt- test was used for within group comparison. Mean, median, SD, and percentiles were
calculated to describe continuous variables. AP-value of ≤ 0.05 was considered significant.

RESULTS
Subjects of all three groups were comparable, baseline MVCs of the legs, and baseline muscle
protein plasma concentrations did not differ significantly between groups before the first
training task. ANOVA and ANCOVA with repeated measures with one grouping and two
within factors showed a significant influence of the additional training regimens on the time
courses of CK, MHC, and MVC. Criterion measures differed significantly between the three
training groups in response to the last training task. Although the subjects of group A and B
performed a high-force eccentric training, no significant differences between the baseline
plasma CK and MHC values before the first (base I) and last (base 2) training task were
found. Cardiac TnI could not be detected in any sample taken, which excluded a protein
release from the heart (cardiac beta-type MHC) and provides evidence for an injury of slow-
twitch skeletal muscle fibers.

Responses to the Initial Training Task

Muscle soreness. Before exercise, all 30 subjects reported no perceived muscle

soreness (rating 0 in all) of the quadriceps muscles. Soreness and pain assessed by
questionnaire were maximal 24-48 h after eccentric exercise and declined on the subsequent
days (mean ratings seeFig. 1).

MVC. Baseline MVC values of the exercised leg of all three groups did not differ
significantly before the first eccentric exercise bout, and one-way ANOVA showed no
significant between-group difference in the observed amount of decrease in MVC after this
first training task as well (seeTable 2).

Muscle protein release. In all subjects baseline CK and MHC concentrations measured
before the first training task were within the reference interval. CK and MHC increased
significantly in response to exercise in all three groups without a significant between group
difference (seeTables 3 and 4).

Responses to the Last Training Task

Muscle soreness. Before the last training task, all 30 subjects reported no perceived muscle
soreness (rating 0 in all) of the quadriceps muscles. The last training task caused muscle
soreness (mean rating 3.9) in most subjects of group C which was not significantly different
from the response after the initial training task. Group A and B subjects, by contrast, did not
complain of any muscle soreness in response to the last training task(see Fig. 1).

MVC. MVC values before the last training task of group B were significantly higher
compared with baseline values before the first training task (base 1). Both other groups
showed no significant increase in MVC. Moreover, MVC values before the last training task
of group B were significantly higher compared with both other groups. After the last (second)
training task subjects of group C again sustained a significant decrease in muscle force
generation (see Table 2). This decrement in the MVC of the loaded leg was not significantly
smaller than that after the first exercise bout. In both other training groups the last training
task did not lead to significant changes in the loaded leg's MVC (seeTable 2).

Muscle protein release. In all subjects baseline CK and MHC concentrations measured
before the last training task did not differ significantly from baseline values before the first
training task (base 1). The last training task did not cause a significant increase in plasma CK
and MHC in subjects of group A and B (see Tables 3 and 4, respectively). The subjects of
group C again showed a significant increase in CK and MHC (see Tables 3 and 4,
respectively). The magnitude of increase in both muscle proteins did not differ significantly
from that after the first training task.

DISCUSSION
We investigated muscular adaptation and strength increase during the early phase of eccentric
heavy-resistance training. The new approach in the present study was to compare the
influence of the frequency of specific standardized training tasks on muscle injury and
adaptation. Maximum isometric voluntary contraction (MVC) force, muscle soreness, and
plasma concentrations of muscle proteins were used as indictors of muscle injury and
adaptation before and after a period of 7 wk. One training regimen (Group A) included a low
training frequency, a second (Group B) included a higher training frequency, and a third
(Group C) no additional training tasks. In all training groups the first bout of heavy eccentric
exercise resulted in a significant temporary loss of the exercised muscles capacity for force
production, muscle soreness, and a significant delayed increase in muscle proteins (CK,
MHC). MHC increase indicates a leakage of the muscle cell membrane and a dissociation or
degradation of the contractile apparatus (6). In the absence of an increase in cTnI, the MHC
increases indicate severe damage to some slow-twitch skeletal muscle fibers. This confirms
prior observations(22,31).
Despite repeated high-force eccentric training in groups A and B, muscle protein plasma
concentrations returned to reference ranges and muscle soreness vanished. In all groups there
was no significant difference between CK, MHC, and muscle sorenessbaseline values before
the first and last training task. After the training period, there was no significant difference in
the response to the last training task of group A and group B. There were no significant
increases in plasma levels of muscle proteins, no muscle soreness, and no temporary loss of
MVC in both groups, which indicates a complete adaptation of the exercised muscles.
Armstrong et al. (1) suggested that the first bout of eccentric exercise damages the most
vulnerable fibers which are then replaced by more stress resistant fibers. However,
Newham(25) suggested that the repeated-bout effect may result from changes in the property
and amount of the connective tissue making the fibers more resistant to subsequent injury.
Kuipers (17)postulated that the adaptation is probably attributed to a change in recruitment and
to an increase in connective tissue as well. The lack of a significant increase in muscle protein
plasma concentrations after the last training task in groups A and B is consistent with all these
different hypotheses.

Only in group B did MVC significantly increase during the training period. The MVC of
group B was also significantly higher compared with both other groups after 6 wk from the
onset of training. This shows that a higher training frequency is necessary to improve MVC,
whereas a low frequency is sufficient to maintain muscle adaptation. One explanation may be
that the initial increase in strength during the early phase of eccentric training is accounted for
by a neuronal adaptation(10,14,28). However, Staron et al.(32) showed that within the first several
weeks of heavy-resistance training changes in the quality of protein without hypertrophy is
the primary adaptation reaction. Whether these increases in strength result from training
induced adaptations of the neuromuscular system or from other factors such as the quality of
muscle proteins cannot be ascertained from our data. It seems unilikely that during the early
phase of eccentric training only a single mechanism is responsible for the observed increase in
strength. Concerning the adaptation process triggered by low frequency of training tasks,
there is only one comparable study, from Balnave and Thompson (2). This study included an
eccentric training once a week for a period of 8 wk with a 40-min walk down with a 25%
gradient on a motor-driven treadmill at 6.4 km·h-1. Similar to our observations, the reactions
to the first and last training tasks were assessed. Balnave and Thompson (2) showed that
eccentric training reduces muscle soreness, the serum muscle protein response, and muscle
function impairment. However, the progressively reduced CK responses still showed a
significant rise after the second exercise test in the training group. This is in contrast to our
data and may be caused by the differences in training regimens as well as the intensities of
eccentric exercise.

Muscular adaptation in our study was exclusively found in groups A and B but not in group
C. This is in contrast to other investigations which used maximal eccentric contractions of the
forearm flexors as an exercise protocol(4) and to the reaction of the control group in the study
of Balnave and Thompson (2). For the control group Balnave and Thompson postulated a long-
lasting adaptation effect based on the fact that recovery in fatigue index response was more
rapid after the second walk. Otherwise there was no significant difference between increases
in serum CK, no significant difference in the fatigue index response, and only serum
myoglobin values increased to a lesser extent after the second down-hill walk. Clarkson et
al. (4) performed one bout of eccentric exercise using the forearm flexor muscles. This
produced a period of adaptation between 6 and 10 wk such that muscle was more resistant to
damage from a subsequent bout of exercise and a reduction in CK response after 6 months (4).
All subjects of group C of our study experienced muscle soreness, showed a decrease in
MVC, and had delayed increases in muscle protein plasma concentrations after the last
(second) training task. There were no significant differences from the reactions after the first
training task. In a previous study (23) with the same eccentric exercise protocol, we observed a
rapid adaptation after 4 and 13 d such that the exercised muscle was more resistant to a
subsequent bout of eccentric exercise. This adaptation was lost after 6 wk without specific
eccentric training. The difference from prior observations of Balnave and Thompson (2) may
be explained by the different eccentric exercise regimens. The main difference is that
downhill walking excludes supramaximal eccentric muscle loads which were used in our
study. If this is the reason for different adaptation periods, a connection between the level of
eccentric muscle loading and the duration of adaptation must be postulated. Regarding long-
lasting muscular adaptation after a single eccentric exercise bout, we could not confirm the
results of Clarkson et al.(4). We and this group used comparable exercise regimens which,
however, involved different muscle groups (m. quadriceps femoris and m. biceps brachii,
respectively). M vastus lateralis and m. biceps brachii do not differ in muscle fiber type
composition (50% slow-twitch and 50% fast-twitch muscle fibers) (9,27), which excludes a
predominant influence of the muscle fiber type composition on the adaptation to eccentric
exercise.

Study limitations. The intensity of eccentric maximal contractions used in the present study,
in particular in the first session, was higher than those applied in earlier studies(2,12) or in the
current typical training program used by athletes or others in the fitness community.
However, we used this approach to be able to better study also the adaptation to the eccentric
damage stress over the early phases of the training program. Furthermore, the use of a
isometric strength testing protocol may be a possible limitation of the study design. Isometric
strength testing was frequently used in previous studies on provoked neuromuscular
adaptation and the efficacy of training regimens (2,11,32). According to more recent results (12), a
specific eccentric strength testing could have been more appropriate because isometric force
production underestimates the benefits of eccentric training and the eccentric force production
ability (12).

In summary, our data indicate that during the early phase of eccentric muscle training a
training task once a week is sufficient to maintain muscle adaptation, whereas training of at
least 2 times a week is needed for strength increases to occur even with the use of
supramaximal eccentric training. The gains during the early phase of training are greater with
the higher frequency than with the low frequency training regimen without causing any
additional muscle damage. The muscular adaptation after a single bout of high-force eccentric
exercise of the thigh muscles was lost after 7 wk without training.

This study was supported in part by the contract 4982 from the Austrian Nationalbank Jubilee
Fond.

Present address of S. Sorichter: Medizinische Universitätsklinik, Abteilung Pneumologie,

Universitätsklinik Freiburg, Hugstetterstr. 55, D-79106 Freiburg, Germany.

Present address of J. Mair, P. Secnik and V. Parrak: Institut für Med. Chemie und Biochemie,
Universität Innsbruck, Fritz-Pregl-Str. 3, A-6020 Innsbruck, Austria.

Present address of A. Kollor: Institut f¨r Sport- und Kreislaufmedizin, Universitätsklinik

Innsbruck, Anichstr. 35, A-6020 Innsbruck, Austria.

Present address of Ch. Haid: Orthopädische Universitätskilinik, Universitätsklinik Innsbruck,

Anichstr. 35, A-6020 Innsbruck, Austria.
Present address of E. Müller: Institut für Sportwissenschaften, Universität Salzburg,
Akademiestr. 26, A-5020 Salzburg, Austria.

Address for correspondence: Prof. Dr. B. Puschendorf, Institut für Medizinische Chemie und
Biochemie, Fritz-Pregl-Str. 3, A-6020 Innsbruck, Austria. E-mail: Johannes.Mair@uibk.ac.at.

Figure 1-Muscle soreness before and after the first and last training tasks. Muscle

soreness and pain before (base) and 1, 2, 3, 4, and 5 d after the first (1) and last (2) eccentric
training task were assessed by questionnaire. On a scale of 0 (normal) to 10 (very sore)
subjects rated the preceived soreness of the quadriceps muscle. Data given as mean. For group
classification see :
Table 1.

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Keywords:
EXERCISE-INDUCED MUSCLE INJURY; MUSCLE SORENESS; MYOSIN HEAVY
CHAIN; CREATINE KINEASE
©1997The American College of Sports Medicine