Eur J Appl Physiol (2010) 108:12471258

DOI 10.1007/s00421-009-1319-8

ORIGINAL ARTICLE

Muscle adaptations and performance enhancements

of soccer training for untrained men
Peter Krustrup Jesper F. Christensen Morten B. Randers Henrik Pedersen
Emil Sundstrup Markus D. Jakobsen Birgitte R. Krustrup Jens J. Nielsen
Charlotte Suetta Lars Nybo Jens Bangsbo

Accepted: 2 December 2009 / Published online: 29 December 2009

Springer-Verlag 2009

Abstract We examined the physical demands of smallsided soccer games in untrained middle-age males and
muscle adaptations and performance effects over 12 weeks
of recreational soccer training in comparison with continuous running. Thirty-eight healthy subjects (2043 years)
were randomized into a soccer (SO), running (RU) and
control (CO) group. Twothree weekly 1-h training
sessions were performed. Muscle lactate (30.1 4.1 vs.
15.6 3.3 mmol/kg d.w.), blood lactate, blood glucose
and time above 90% HRmax (20 4% vs. 1 1%) were
higher (p \ 0.05) during training in SO than in RU. After
12 weeks of training, quadriceps muscle mass and mean
muscle fibre area were 9 and 15% larger (p \ 0.05) in SO,
but unaltered in RU, and in SO, the fraction of FTx fibres
was lowered (10.7 1.8 vs. 17.9 3.2%). In SO, citrate
synthase activity was 10 and 14% higher (p \ 0.05) after 4
and 12 weeks, but unaltered in RU. After 4 weeks VO2max
and Yo-Yo IE2 performance were elevated (p \ 0.05) to a
similar extent in SO (7 and 37%) and RU (6 and 36%) but
increased further (p \ 0.05) from 4 to 12 weeks in SO (6
and 23%). In SO, 30-m sprint performance was improved
(p \ 0.05) by 0.11 0.02 s. Blood lactate during running
Communicated by Susan Ward.
P. Krustrup (&)
J. F. Christensen
M. B. Randers

H. Pedersen
E. Sundstrup
M. D. Jakobsen

B. R. Krustrup
J. J. Nielsen
L. Nybo
J. Bangsbo
Department of Exercise and Sport Sciences, Section of Human
Physiology, Copenhagen Muscle Research Centre,
University of Copenhagen, The August Krogh Building,
Universitetsparken 13, Copenhagen 2100, Denmark
e-mail: pkrustrup@ifi.ku.dk
E. Sundstrup
M. D. Jakobsen
C. Suetta
Institute of Sports Medicine Copenhagen,
Bispebjerg University Hospital, Copenhagen, Denmark

at 11 km/h was lowered (p \ 0.05) from 0 to 4 and 4 to

12 weeks (2.6 0.3 vs. 3.8 0.6 vs. 6.1 0.9 mM) and
from 0 to 12 weeks in RU. No changes occurred for CO. In
conclusion, recreational soccer organized as small-sided
games stimulates both aerobic and anaerobic energy turnover and is an effective type of training leading to significant cardiovascular and muscular adaptations as well as
performance enhancements throughout a 12-week training
period.
Keywords Muscle metabolites
Blood lactate

Activity profile
Enzyme activity

Respiratory exchange ratio
Fibre size

Sprint performance
Yo-Yo intermittent endurance test

Football

Introduction
Soccer practice is physically demanding for experienced
players. Thus, for elite and sub-elite players multiple
intense activities are performed during small-sided games
and drills with reduced pitch size and number of players,
resulting in significantly elevated heart rate and blood
lactate (Bangsbo 1998; Coutts et al. 2009; Impellizzeri
et al. 2006; Jones and Drust 2007; Little and Williams
2006, 2007; Owen et al. 2004). However, the activity
pattern and physical response of non-experienced soccer
players during such small-sided games, 5v5 or 7v7, have
not been evaluated. Furthermore, muscle biopsies have not
been collected for players during this type of soccer
training to determine muscle substrate utilization and
anaerobic energy turnover.
A number of studies have shown improvements in
maximal oxygen uptake (VO2max) and intermittent exercise

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performance of experienced soccer players after a period

of soccer training (Bangsbo 1994; Impellizzeri et al.
2006; Krustrup et al. 2003, 2006a; McMillan et al. 2005).
In addition, the effect of performing additional smallsided games with high physiological strain has been
compared with the effect of carrying out supplementary
running interval training sessions (Impellizzeri et al.
2006). In the study by Impellizzeri et al. (2006) players
were followed in the pre-season period, and twice a week
part of their normal training was devoted to small-sided
games selected to raise the heart rate to 9095% of
HRmax. The effect was compared with interval running
consisting of four bouts of exercise lasting 4 min with
3 min active recovery. Similar improvements in VO2max and
continuous exercise performance were observed. However,
the effect of soccer training on intermittent exercise
performance was not studied and only a limited number of
studies have investigated changes in lean body mass and
muscle adaptations, such as muscle fibre distribution, fibre
area and enzymes after a period of soccer training
(Bangsbo and Mizuno 1988). Furthermore, no studies have
focused on changes in muscle strength and balance as a
consequence of ordinary soccer training and the effect of
regular soccer training for untrained subjects has not been
examined.
Thus, the purpose of the present study was to examine
the physical demands of small-sided games in untrained
subjects and compare this to running training of the same
duration. An additional purpose was to study the time
course of muscle adaptation and effect on performance
during various type of exercise in a period of soccer
training and compare that to a similar period of running
training in untrained middle-aged men.

Eur J Appl Physiol (2010) 108:12471258

approved by the local ethical committee of Copenhagen

(14606; HC-2007-0012).
Design
The subjects were matched and randomly assigned to a
soccer group (SO, n = 14), a running group (RU, n = 13)
or a control group performing no physical training (CO,
n = 11). One soccer player, two runners and one control
subject dropped out due to injuries occurring during
training or testing. In addition, one soccer player and one
runner performed too few tests in the last testing round to
be included in the analysis. The six subjects who did not
complete the study did not differ in age, body composition
or training status from the rest of the subjects. For the
participants who completed the study (SO, n = 12; RU,
n = 10; CO, n = 10) no group differences were present in
pre-intervention values for age, body mass, fat percentage
and VO2max. The participant in SO and RU carried out a
12-week soccer and running training programme, respectively, whereas the participants in CO continued their daily
life activities during the period. The subjects were examined before the intervention period as well as 4 and
12 weeks into the intervention period in order to examine
the muscular and cardiovascular adaptations to the training
and to investigate possible changes in exercise performance. The testing protocol included muscle biopsies,
blood samples, DXA scans, treadmill tests and a series of
performance tests (see below). In addition, video filming,
muscle biopsy sampling and collection of blood samples
were performed for both training groups during a training
session after 4 weeks in order to investigate the activity
profile and the physiological response to recreational soccer and running.

Methods

Training intervention

Subjects

Outdoor training was performed twothree times a week

for 12 weeks, as described in Krustrup et al. 2009. Briefly,
the soccer sessions consisted of ordinary five-a-side and
seven-a-side matches on a 3040 m wide and 4060 m
long natural grass pitch. The running sessions consisted of
continuous moderate intensity running at 82% of individual
maximal heart rate. The duration of the training sessions
was 60 min for both training groups. The total number of
training sessions were 27.8 (2.3 per week) and 30.4 (2.5 per
week) for the participants in SO and RU, respectively.

Thirty-eight healthy untrained males aged 2043 years

(body mass, fat percentage and VO2max 84.4 2.4
(SEM) kg, 24.5 1.1% and 39.4 0.9 ml min-1 kg-1,
respectively) volunteered to participate in the study. Participants were recruited through adverts in local newspapers, and flyers posted in educational departments and
other public institutions in Copenhagen. None of the participants were on any medication and none were smokers.
None of the subjects had been involved in regular physical
activity for at least 2 years. The subjects were fully
informed of the risks and discomforts associated with the
experimental procedures, and all provided written consent.
The study was carried out in accordance with the guidelines contained in the Declaration of Helsinki and was

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Measuring and test procedures

Subjects were familiarized to all test procedures on 13
occasions. No physical activities were performed 2 days
prior to testing. For the training groups, the 4- and

Eur J Appl Physiol (2010) 108:12471258

12-week-muscle biopsies were obtained 4252 h after a

training session.
Muscle biopsies
Muscle biopsies were obtained from m. vastus lateralis
under local anaesthesia using the Bergstrom technique. For
the training groups, a total of eight muscle biopsies were
collected, i.e. three prior to the intervention, two after
4 weeks of training and three after 12 weeks of training.
One of the muscle biopsies for each time point was a
resting biopsy whereas the others were exercise/recovery
biopsies (see below). The control subjects had resting
biopsies taken prior to and after the 12-week-intervention
period.
Sub-maximal and maximal treadmill testing
After 0, 4 and 12 weeks (0 and 12 weeks for CO) pulmonary gas exchange (CPX MedGraphics, USA), heart
rate (Polar Team System, Polar Electro Oy, Kempele,
Finland) and arm vein blood sampling were performed
during a standardized treadmill test with 6-min bouts of
walking at 6.5 km h-1 and sub-maximal running at 8.0, 9.5
and 11.0 km h-1, interspersed with 2-min rest periods.
Respiratory exchange ratio (RER) was calculated over the
last 3 min of running at each speed, as the carbon dioxide
release divided by oxygen uptake, and used as an indicator
of fat oxidation during sub-maximal exercise. After a
15-min rest period, the subjects carried out an incremental
test to exhaustion, consisting of 4 min of running at
9.5 km h-1 followed by stepwise 1 km h-1 speed increments
each 30 s until exhaustion.
Muscle mass and fat percentage
After 0, 4 and 12 weeks (0 and 12 weeks for CO) quadriceps muscle mass was determined by anthropometrical
measurements as described by Krustrup et al. (2004). Prior
to and 2 days after (i.e. 0 and 12 weeks) fat, muscle and
bone mass was determined by whole body DXA scans
(LUNAR, GE Medical Systems, USA) from 8 to 10 a.m.
under standardized conditions after an overnight fast.

1249

running speed as well as at exhaustion and 1, 3, 5, 10 and

15 min into the recovery period. Blood samples during the
test were collected with the subjects standing upright during the 5-s recovery periods, and blood samples in the posttest recovery period were collected with the subject lying
down in a supine position. Muscle samples were collected
from m. vastus lateralis at rest, at exhaustion and 3 min
into the recovery period as described in Mohr et al. (2007).
The subjects in SO and RU also performed a 30-m sprint
test after 0 and 12 weeks using infra-red timing gates with
a precision of 0.01 s (Time It, Eleiko Sport, Halmstad,
Sweden). Each participant had two trials separated by
5 min of rest and the best time was used as the test result.
Balance was determined by a 1-min single-leg Flamingo
balance test (Adam et al. 1988; Deforche et al. 2003).
Maximal isometric quadriceps and hamstring contractions
were performed during static knee extensions/flexions at
a knee joint angle of 70 (0 = full knee extension)
(Aagaard et al. 2002). The measurements were exerted
in an isokinetic dynamometer (KinCom Coummicator,
Chattecx, Chattanooga, TN, USA).
Measurements during training
Heart rate was determined during all training sessions.
During a single training session 4 weeks into the intervention period, blood samples were collected by 2 ml
syringes from a catheter placed in an antecubital arm vein
for measurements of blood lactate and glucose. Samples
were collected at rest, after 5, 15, 30, 45 and 60 min of
training as well as 15 and 30 min into the recovery period.
In addition, muscle biopsies were collected from muscle
vastus lateralis at rest and during the last 15 min of training
(i.e. 4560 min). During training sessions after 4 and
6 weeks, a total of nine players were video-filmed individually close up during the entire training session. VHS
movie cameras (NV-M50, Panasonic, Germany) were
positioned at the side of the pitch, at the level of the
midfield line, at a height of about 6 m and at a distance of
30 m from the touchline. The videotapes were later
replayed on a monitor for computerized coding of the
activity pattern.
Activity profile analyses

Performance testing
After 0, 4 and 12 weeks (0 and 12 weeks for CO) the
participants carried out a Yo-Yo intermittent endurance
level 2 test (Bangsbo 1995). Heart rate was recorded
throughout the test and during the first 15 min of recovery.
After 0 and 12 weeks for SO and RU, blood samples and
muscle biopsies were also collected. Blood samples were
collected at rest, after the 5-min warm-up period, at each

The players activity profile was assessed by a video based

time-motion analysis system (Bangsbo et al. 1991;
Krustrup et al. 2005). The number of occurrences and
the duration of each occurrence were noted for the following locomotor activities: standing (0 km h-1), walking
(4 km h-1), jogging (8 km h-1), low-speed running
(12 km h-1), moderate-speed running (15 km h-1), highspeed running (18 km h-1), sprinting (25 km h-1) and

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sideways/backward running (10 km h-1). The activities

were later divided into four categories expressing the
exercise intensity: (1) standing; (2) walking; (3) lowintensity running, encompassing jogging, low-speed running and sideways/backward running and (4) high-intensity
running, consisting of moderate-speed running, high-speed
running and sprinting.
Muscle analyses
A part of the muscle sample (*60 mg wet weight) was
immediately frozen in liquid N2 and stored at -80C. The
frozen sample was weighed before and after freeze-drying
to determine water content. After freeze-drying, the muscle
samples were dissected free of blood, fat and connective
tissue and about *1 mg dry wt tissue was extracted in a
solution of 0.6 M perchloric acid (PCA) and 1 mM EDTA,
neutralized to pH 7.0 with 2.2 M KHCO3 and stored at
-80C until analyzed for lactate and creatine phosphate
(CP) by a flurometric assay (Lowry and Passonneau
1971). Muscle pH was measured by a small glass electrode (Radiometer GK2801, Copenhagen, Denmark) after
homogenizing a freeze-dried muscle sample of about
2 mg d.w. in a non-buffering solution containing 145 mM
KCl, 10 mM NaCl and 5 mM iodoacetic acid. Another
12 mg dry wt muscle tissue was extracted in 1 M HCl
and hydrolyzed at 100C for 3 h and the glycogen content
was determined by the hexokinase method (Lowry and
Passonneau 1971). In addition, 2 mg dry wt muscle tissue
was analyzed for citrate synthase (CS) and 3-hydroxyacyl-CoA dehydrogenase (HAD) activity using fluourometric methods with NAD-NADH coupled reactions
(Lowry and Passonneau 1971). The remainder of the
muscle tissue was mounted in an embedding medium
(OCT Tissue-Tek, Sakura Finetek, Zoeterwoude, The
Netherlands) frozen in pre-cooled isopentane and analyzed histochemically for fibre area and capillaries
(Brooke and Kaiser 1970).

Eur J Appl Physiol (2010) 108:12471258

intervention within-group differences in physiological

response to Yo-Yo IE2 testing and treadmill testing were
also evaluated by a two-way ANOVA exercise time/running speed 9 intervention). Significant main effects were
subsequently analyzed using a Tukeys post hoc test. Preintervention and intervention-induced differences in absolute values for performance and muscular variables as well
as between-group differences in pre-post intervention delta
values were evaluated by simple ANOVA tests. When a
significant main effect was detected data were subsequently analyzed using a Tukeys post hoc test. Normality
was verified using the ShapiroWilk W test. A significance
level of 0.05 was chosen. Data are presented as mean
standard error of the mean (SEM).

Results
Activity profile during training
The total number of activity changes over a 1-h soccer
training session was 886 44, corresponding to a change
in activity every *4 s (Fig. 1). A total of 16 1 sprints
and 98 5 high-intensity runs ([15 km/h) were performed
with a mean duration of 1.9 0.1 and 2.3 0.1 s,
respectively. The number of backwards/sideways runs was
59 10 with a mean duration of 2.5 0.2 s. Total distance covered during a soccer training session was
5.00 0.09 km of which 0.88 0.04 and 0.22 0.02 km
were covered with high intensity running and sprinting,
respectively. During training the soccer players stood still,
walked or ran at low intensity for 20.8 1.8, 51.9 1.1
and 22.1 1.7% of the time, respectively, whereas they
performed high-intensity running for 5.1 0.2% of the
time. The number of headers and tackles performed during
a training session were 1.8 0.6 and 11.3 2.0,
respectively.
Physiological response to training

Blood analyses
Blood samples were obtained in 2 ml syringes without
heparin. Within 10 s of sampling 100 ll of blood was
hemolyzed in an ice-cold 100 ll Triton X-100 buffer
solution, and was later analyzed for lactate and glucose
using an YSI 2300 lactate analyser (Yellow Spring
Instruments, Yellow Springs, OH, USA).
Statistics
Between-group differences in the metabolite response to a
training session were evaluated by a two-way analyses of
variance (ANOVA; exercise time x group). Pre-post

123

Blood lactate was higher (p \ 0.05) in SO than in RU at

several time-points during a training session (30 min:
5.2 0.6 vs. 3.7 0.4 mM) (Fig. 2a). Peak blood lactate
during training was also higher (p \ 0.05) for SO than RU
(5.9 0.6 vs. 4.3 0.4 mM). Blood glucose was higher
(p \ 0.05) in SO than in RU throughout a training session
(30 min: 5.5 0.5 vs. 3.3 0.4 mM) (Fig. 2b). The
lowest blood glucose value during training was also higher
(p \ 0.05) in SO when compared with RU (3.7 0.2 vs.
2.8 0.2 mM).
At the end of the training session, muscle lactate was
higher (p \ 0.05) in SO than in RU (30.1 4.1 vs.
15.6 3.3 mmol kg-1 d.w.; Table 1). Muscle CP in SO

Eur J Appl Physiol (2010) 108:12471258

1251
Table 1 Muscle metabolites (mmol kg-1 d.w.) and after a soccer
training session (SO, n = 11) and a running training session (RU,
n = 8) performed 4 weeks into the 12-week training period

350

Number of activities

300

SO

250

Before

After

Lactate

11.6 2.7

30.1 4.1*,#

9.7 2.1

CP

80.5 8.0

50.3 5.0*

85.2 7.9

59.4 10.4*

Glycogen

422 28

304 31*

402 19

303 26*

200
150

RU
Before

After
15.6 3.3

100

Mean SEM are presented

* Significant difference from before training

50

Significant difference from RU

0
St

Ls

Ms

Hs

Sp

HI

Activity

Fig. 1 Activity profile during a 1-h soccer training session, expressed

as the number of occurrences of various locomotor modes including
standing (St), walking (W), jogging (J), low speed running (Ls),
moderate speed running (Ms), high speed running (Hs), sprinting (Sp),
sideways/backward running (B), as well as the total amount of high
intensity running (HI, sum of Ms, Hs and Sp). Mean SEM (n=9) are
presented

SO
RU

Blood lactate (mmol/l)

6
#

Peak
values

5
4

and RU was 37 and 30%, respectively, lower (p \ 0.05)

when compared with before the training session (Table 1).
Muscle glycogen decreased (p \ 0.05) by 118 27 and
100 33 mmol kg-1 d.w. in SO and RU during the
training session, respectively, with no difference between
groups (Table 1).
The average heart rate during training was 166 3 and
159 4 b.p.m. in SO and RU, which corresponded to
82 2 and 82 1% of HRmax for SO (202 2 b.p.m.)
and RU (195 3 b.p.m.), respectively. The time above
90% of HRmax was higher (p \ 0.05) in SO than RU
(20 4 vs. 1 1%). No changes were observed in average heart rate or time above 90% HRmax from the first to
the last week of training.
Effects of the 12-week training period

3
2

Muscle adaptations

1
0
0

Blood glucose (mmol/l)

10

20

30

40

50

60

70

6.0
5.5

90

SO
RU

80

5.0

4.5
4.0

3.5

Lowest
values

3.0
2.5
0.0
0

10

20

30

40

50

60

70

80

90

Time (min)

Fig. 2 Blood lactate (a) and blood glucose (b) before, during and
after a soccer (SO, n = 11) and a running training session (RU,
n = 8) performed 4 weeks into the 12-week training period.
Mean SEM are presented. #Significant difference from RU

In SO, the mean muscle fibre area was 15% larger

(p \ 0.05) after compared with before the 12-week training
period (5,546 240 vs. 4,828 233 lm2), whereas no
significant changes occurred for RU and CO (Fig. 3).
Estimated quadriceps muscle mass was 9% higher
(p \ 0.05) in SO after 12 weeks of training (2.25 0.08
vs. 2.07 0.07 kg), whereas no changes were observed
for RU (2.30 0.14 vs. 2.22 0.12 kg) and CO
(2.32 0.09 vs. 2.28 0.10 kg). In SO, the fraction of
FTx fibres was 10.7 1.8% after 12 weeks of training,
which was lower (p \ 0.05) than before training
(17.9 3.2%), whereas fibre type distribution was not
significantly altered in RU or CO (Table 2).
After 12 weeks of training the number of capillaries per
fibre was 22 and 16% higher (p \ 0.05) in SO and RU,
respectively (Table 2). When capillarisation was expressed
as capillaries per area, the changes over 12 weeks did not
reach significance (12% for SO and 10% for RU). For CO,
no changes occurred in capillarisation over the 12 weeks
(Table 2).

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Eur J Appl Physiol (2010) 108:12471258

Citrate synthase activity (mol/g d.w./min)

0 wks
4 wks
12 wks

50
45
40

*+

35
30
25
0

SO

Mean muscle fibre size (m )

RU

CO

6250
6000

0 wks
4 wks
12 wks

*+

5750
5500
5250
5000
4750
4500
0

SO

RU

CO

Fig. 3 Muscle citrate synthase activity (a) and fibre size (b) before as
well as after 4 and 12 weeks of soccer practice (SO, n = 12) and
running (RU, n = 10) for untrained men, or continuation of an
inactive life-style (CO = 10). Mean SEM are presented. * Significant difference from 0 weeks. ?Significant difference from CO

In SO, citrate synthase activity was 10 and 14% higher

(p \ 0.05) after 4 and 12 weeks of training (39.3 2.0
and 40.7 2.3 vs. 35.6 2.2 lmol g-1d.w. min-1), but
unaltered in RU and CO (Fig. 3). HAD activity was not
significantly altered after 12 weeks of training for neither
SO, RU or CO (Table 2).
Physiological and functional capacity
After 12 weeks of training, 30-m sprint time was
0.11 0.02 s better (p \ 0.05) for SO (4.64 0.05 vs.
4.75 0.06 s), but unaltered for RU (4.77 0.04 vs.
4.76 0.05 s). In SO, maximal isometric hamstring
strength (normalized to body weight) was elevated
(p \ 0.05) over 12 weeks by 11% (1.42 0.09 vs. 1.28
0.06 Nm kg-1), whereas maximal isometric quadriceps
strength was not significantly altered (3.10 0.11 vs.
3.14 0.08 Nm kg-1). No changes occurred for RU and

123

CO neither for maximal isometric hamstring strength

(1.34 0.05 vs. 1.36 0.05 and 1.44 0.09 vs.
1.41 0.09 Nm kg-1, respectively) nor for isometric
quadriceps strength (2.88 0.17 vs. 2.83 0.19 and
3.09 0.14 vs. 2.95 0.15 Nm kg-1, respectively).
After 12 weeks of training, the number of falls in the onelegged 1-min Flamingo balance test was lower (p \ 0.05)
both in SO (7 1 vs. 13 2) and RU (10 1 vs.
17 2) whereas it was unchanged in CO (13 2 vs.
14 3).
Yo-Yo IE2 performance before the training period was
655 37, 569 85 and 662 80 m for SO, RU and CO,
respectively. In SO, Yo-Yo IE2 performance was elevated
(p \ 0.05) by 235 44 m after 4 weeks and further after
12 weeks (420 27 m, p \ 0.05), whereas in RU performance was enhanced (p \ 0.05) by 187 33 m after
4 weeks with no further increase from 4 to 12 weeks
(Fig. 4). Yo-Yo IE2 performance was not significantly
altered in CO (Fig. 4). In SO, time to exhaustion in
the incremental treadmill test was 17% higher (p \ 0.05)
after 4 weeks and was further elevated (p \ 0.05) by 8%
after 12 weeks (8.38 0.16 and 7.82 0.11 min vs.
6.69 0.15 min). In RU, time to exhaustion was elevated
(p \ 0.05) by 20% after 4 weeks and further (p \ 0.05) by
6% from 4 to 12 weeks (7.99 0.37 and 7.57 0.23 min
vs. 6.30 0.28 min). In CO, time to exhaustion was
6% higher (p \ 0.05) after 12 weeks (7.68 0.28 vs.
7.26 0.31 min).
In SO, maximal oxygen uptake was 39.6 1.5
ml min-1 kg-1 before training and was elevated by 7%
after 4 weeks and further by 6% during the following
8 weeks of training (Fig. 4). In RU, the maximal oxygen
uptake was 39.3 2.1 ml min-1 kg-1 before training and
increased (p \ 0.05) by 6% over the first 4 weeks, with no
further change after 12 weeks (Fig. 4). After 12 weeks of
training, peak ventilation was 17 3 and 14 3 l min-1
higher (p \ 0.05) in SO and RU, respectively, than before
the training period. In CO, no changes occurred in peak
ventilation (1 3 l min-1) and maximal oxygen uptake
(Fig. 4).
Body mass was lowered (p \ 0.05) by 1.1 0.2 and
1.0 0.3 kg, for SO and RU, respectively, over 12 weeks,
with no change for CO. Lean body mass was 1.7 0.4 kg
higher (p \ 0.05) in SO after the 12 weeks, whereas no
significant changes occurred for RU (0.6 0.3 kg) and
CO (0.3 0.4 kg).
Physiological response to walking and sub-maximal
running
In SO, heart rate during walking at 6.5 km h-1 and running
at 8.0, 9.5 and 11.0 km h-1 was lowered (p \ 0.05) by
1017 b.p.m. after 4 weeks and by 1322 b.p.m. after

Eur J Appl Physiol (2010) 108:12471258

1253

Table 2 Muscular metabolites, enzyme activity, fibre type distribution and fibre sizes after 0, 4 and 12 weeks of soccer practice (SO, n = 12)
and running (RU, n = 10) for untrained men, as well as after 0 and 12 weeks for an inactive control group (CO, n = 10)
SO
0 weeks

RU
4 weeks

12 weeks

CO

0 weeks

4 weeks

12 weeks

0 weeks

12 weeks

Glycogen content
(mmol/kg d.w.)

464 37

422 28

470 31

412 29

402 19

494 45

374 43

376 32

CP content
(mmol/kg d.w.)

89.2 6.8

80.5 8.0

85.9 10.5

82.7 9.4

85.2 7.9

77.6 13.9

92.0 4.7

81.5 9.0

CS activity (lmol/
g d.w./min)

35.6 2.2

39.3 2.0*

40.7 2.3*

33.0 2.8

35.6 2.2

35.4 2.3

42.1 3.2

37.5 4.4

HAD activity
(lmol/g d.w./min)

25.4 2.3

27.2 1.8

26.6 1.5

28.1 1.9

29.2 2.3

29.6 2.4

31.0 1.5

27.6 2.3

Capillaries
(cap/fibre)

1.76 0.11

2.01 0.12

2.15 0.12*

1.83 0.10

2.02 0.12

2.09 0.12*

2.18 0.08

2.29 0.05

Capillaries
(cap/mm2)

354 25

386 28

395 18

367 19

371 15

404 30

430 24

454 32

ST fibre size
(lm2)

4,711 304 5,417 447 5,683 463

5,229 309 5,490 371 5,436 501

5,056 315

4,733 339

FTa fibre size

(lm2)

5,172 271 5,911 420 5,604 207

5,169 230 5,858 303 5,695 385

4,992 263

4,728 358

FTx fibre size

(lm2)

4,509 336 4,777 419 5,121 317

4,721 285 4,915 465 5,111 502

3,456 1,020 4,484 213

Mean fibre size

(lm2)

4,828 233 5,513 373 5,546 240* 5,075 239 5,507 287 5,367 473

5,065 271

4,767 233

Fibre type
distribution
(% ST fibres)

43.0 3.8

45.3 6.0

47.4 4.7

44.6 3.8

48.4 3.7

47.6 5.9

43.9 5.3

49.0 5.0

Fibre type
distribution
(% FTa fibres)

39.1 2.7

38.1 3.5

41.9 3.9

38.7 2.7

37.5 4.6

40.1 4.3

44.6 3.9

39.9 3.6

Fibre type
distribution
(% FTx fibres)

17.9 3.2

16.5 3.9

10.7 1.8*

16.7 3.9

14.1 3.6

12.2 4.6

11.5 3.0

10.7 1.8

Mean SEM are presented

* Significant difference from 0 weeks

12 weeks of training, respectively, with corresponding

values for RU being 1517 b.p.m. and 1423 b.p.m.,
respectively, whereas no change occurred for CO
(Table 3).
In SO, blood lactate was lower (p \ 0.05) at 9.5 and
11.0 km h-1 after 4 weeks of training with a further
change (p \ 0.05) after 12 weeks (Fig. 5). Over 12 weeks
of training in SO, blood lactate was also lowered
(p \ 0.05) at 6.5 and 8.0 km h-1 (Fig. 5). In RU, after
4 weeks of training blood lactate was unaltered at all
intensities, whereas it was lower (p \ 0.05) at 9.5 and
11 km h-1 after 12 weeks of training (Fig. 5). Blood lactate was unaltered in CO over 12 weeks for all intensities.
In SO, the RER was lower (p \ 0.05) at 8.0 and
9.5 km h-1 after 4 weeks and at 8.0, 9.5 and 11.0 km h-1
after 12 weeks, whereas no significant changes occurred
for RU or CO (Table 3).

After 12 weeks of training in SO, the slow component of

oxygen uptake was lowered (p \ 0.05) by 0.19 l min-1
during running at 9.5 km h-1, respectively, and by
0.08 l min-1 at 11.0 km h-1 and also RU had a lower
(p \ 0.05) slow component at 11.0 km h-1 after 12 weeks
(Table 3). Oxygen uptake after 2 min of running at 8.0, 9.5
and 11.0 km h-1 was not different before and after the
training period for neither SO, RU or CON, whereas oxygen uptake values after 6 min of running at 8.0, 9.5 and
11.0 km h-1 was 0.170.22 l min-1 lower (p \ 0.05)
in SO and 0.130.23 l min-1 lower (p \ 0.05) in RU
(Table 3).
Physiological response to intermittent exercise (Yo-Yo IE2)
Metabolite changes during and in recovery from Yo-Yo
IE2 performed before and after the training period are

123

1254

Maximal oxygen uptake (ml/min/kg)

Eur J Appl Physiol (2010) 108:12471258

Discussion

48
46
44

0 wks
4 wks
12 wks

+
+
* *

42
40
38
36
0
SO

Yo-Yo IE2 performance (m)

1200
1000

RU

CO

* # +
*

800

0 wks
4 wks
12 wks

* *

600
400
200
0
SO

RU

CO

Fig. 4 Maximal oxygen uptake (a) and Yo-Yo Intermittent Endurance level 2 test performance (b; Yo-Yo IE2) before as well as after 4
and 12 weeks of soccer practice (SO, n = 12) and running (RU,
n = 10) for untrained men, or continuation of an inactive life-style
(CO = 10). Mean SEM are presented. * Significant difference
from 0 weeks. Significant difference from 4 weeks. #Significant
difference from RU. ?Significant difference from CO

presented in Fig. 6. The net rate of muscle lactate accumulation during Yo-Yo IE2 was lowered (p \ 0.05) in SO
after 12 weeks (8 1 vs. 17 2 mmol kg-1 d.w. min-1)
and tended to be lower (p = 0.09) in RU (9 2 vs.
18 5 mmol kg-1 d.w. min-1). Similarly, the net rate of
muscle CP degradation was lowered (p \ 0.05) in SO
(9 1 vs. 18 2 mmol kg-1 d.w. min-1) with no difference in RU. After 12 weeks, muscle lactate concentration in SO was lower (p \ 0.05) after 3 min of recovery
(35 4 vs. 58 5 mmol kg-1 d.w.) when compared with
before the training period. After 12 weeks of training the
net rate of muscle H? accumulation during Yo-Yo IE2 was
lowered (p \ 0.05) both in SO and RU.
For SO, heart rate was reduced (p \ 0.05) by 7
12 b.p.m. from 1.5 to 4 min in Yo-Yo IE2 and by 625
b.p.m. from 1 to 15 min of recovery, whereas no significant
changes occurred for RU.

123

The present study examined the effect of a 12-week period

of soccer training and compared it to a similar period with
running training in untrained middle-aged men. Both types
of training lead to improvements in VO2max and Yo-Yo
intermittent recovery test performance after 4 weeks,
whereas only the soccer training group had further
increases during the following 8 weeks. In addition, only
the soccer group had an increase in the quadriceps muscle
mass, mean fibre area, lean body mass as well as muscle
strength after the training period and a reduction in the
relative number of FTx fibres. Apparently, soccer training
in comparison to running provides additional and continuous stimuli for physiological adaptations and performance
development. The average heart rate during training was
the same in the two groups, but the soccer players spent
significantly more time in the heart rate zone above 90% of
HRmax and had a higher accumulation of muscle and blood
lactate during training.
Both the soccer and running group had significant
increases in VO2max during the first 4 weeks of training,
but only the soccer group had a further increase during the
following 8 weeks, indicating that the soccer group maintained the stimuli for cardiovascular adaptations throughout the entire training period. Both groups had a decrease
in heart rate during submaximal continuous running which
is common finding after a period of endurance training
(Beneke and Hutler 2005), but only the soccer group had a
lowering of heart rate during intermittent exercise. These
findings suggest that the type of training is influencing the
cardiovascular responses to various types of exercise. This
also indicates that the adaptations are not only occurring in
central factors like increased blood volume and elevated
ventricular volume (Goodman et al. 2005) but are also
affected by peripheral variables. In accordance, the
improvement in the Yo-Yo intermittent exercise performance after 4 weeks was similar between the two groups,
but only the soccer group increased their performance level
further as a result of the following 8 weeks of training and
after 12 weeks the soccer group performed better than the
runners. In addition, the two training groups improved
continuous incremental exercise performance to a similar
extent, whereas only the soccer group improved sprint
performance. Taken together, the data suggest that the
soccer training is more effective than running in improving
overall exercise performance, and that, in contrast to running, further improvements can be obtained by maintaining
regular soccer training. Impellizzeri et al. (2006) found
similar improvements in VO2max and continuous exercise
performance when the effect of performing additional
small-sided games was compared with supplementary
interval running sessions for experienced soccer players.

Eur J Appl Physiol (2010) 108:12471258

1255

Table 3 Heart rate, pulmonary oxygen uptake after 2 min of

exercise, slow component of oxygen uptake (62 min) and respiratory
exchange ratio (RER) during treadmill walking (6.5 km h-1) and
running (8.0, 9.5 and 11.0 km h-1) before and after 4 and 12 weeks
SO
0 weeks
Heart rate
(b.p.m)

VO2 (l/min)

Slow component
(l/min)

RER

of soccer practice (SO, n = 12) and running (RU, n = 10) for

untrained men, as well as after 0 and 12 weeks for an inactive control
group (CO, n = 10)
RU

4 weeks

12 weeks

0 weeks

CO
4 weeks

12 weeks

0 weeks

12 weeks

6.5 km/h

122 3

112 2*

109 2*

121 6

106 5*

107 5*

115 3

109 4

8.0 km/h

158 5

144 3*

139 3*

150 5

137 5*

134 5*

153 4

145 6

9.5 km/h

176 3

159 3*

155 3*

167 5

151 6*

146 6*

162 6

157 6

11.0 km/h

185 2

167 2*

163 1*

174 5

157 5*

151 5*

172 5

164 4

6.5 km/h

1.40 0.05

1.38 0.05

1.31 0.04

1.40 0.10

1.33 0.11

1.30 0.11

1.44 0.08

1.47 0.10

8.0 km/h

2.17 0.09

2.17 0.12

2.08 0.08

2.21 0.12

2.15 0.13

2.12 0.14

2.14 0.11

2.05 0.10

9.5 km/h

2.40 0.08

2.51 0.13

2.39 0.10

2.51 0.13

2.43 0.16

2.42 0.15

2.41 0.12

2.34 0.10

11.0 km/h

2.71 0.07

2.64 0.14

2.61 0.06

2.64 0.14

2.54 0.15

2.49 0.15

2.77 0.12

2.67 0.13

Peak

3.21 0.09

3.43 0.13*

3.56 0.11*

3.27 0.10

3.47 0.12*

3.54 0.14*

3.54 0.15

3.46 0.13

6.5 km/h

0.03 0.03

0.01 0.03

0.01 0.02

0.05 0.04

0.00 0.03

0.01 0.02

0.02 0.03

0.04 0.04

8.0 km/h

0.12 0.04

0.08 0.06

0.06 0.05

0.13 0.04

0.08 0.05

0.09 0.03

0.10 0.04

0.10 0.02

9.5 km/h

0.22 0.05

0.10 0.04

0.03 0.02*

0.15 0.04

0.14 0.04

0.07 0.04

0.15 0.04

0.16 0.05

11.0 km/h

0.15 0.02

0.16 0.03

0.07 0.02*

0.15 0.02

0.09 0.04

0.07 0.03*

0.13 0.05

0.17 0.04

6.5 km/h

0.91 0.02

0.84 0.02

0.88 0.03

0.88 0.03

0.85 0.03

0.88 0.03

0.86 0.03

0.93 0.02
0.99 0.03

8.0 km/h

0.98 0.02

0.92 0.02*

0.92 0.03*

0.96 0.03

0.91 0.03

0.91 0.02

0.95 0.03

9.5 km/h

1.00 0.02

0.93 0.02*

0.93 0.02*

0.97 0.03

0.92 0.03

0.93 0.02

0.96 0.03

0.93 0.09

11.0 km/h

1.02 0.03

0.97 0.03

0.93 0.03*

0.95 0.05

0.90 0.04

0.91 0.03

0.99 0.02

1.02 0.03

Peak

1.23 0.04

1.25 0.04

1.24 0.03

1.15 0.04

1.20 0.04

1.25 0.03

1.25 0.05

1.27 0.04

Mean SEM are presented

* Significant difference from 0 weeks

However, it is difficult to compare the studies as Impellizzeri et al. (2006) studied experienced players in the preseason where the additional training only constituted a
minimal part of the entire training and the intensity during
the running training was higher than in the present study.
Furthermore, no measurements were made during the
training period. The present study does, however, suggests
that carrying out small-sided games is an effective performance improving type of training. In accordance Jensen
et al. (2009) observed that one additional 30-min aerobic
high intensity training session a week consisting of smallsided drills improved the intermittent exercise performance
of experienced players.
In the present study, the physiological response during
exercise and the muscle adaptations after a period of soccer
training were also investigated. The soccer training elevated the CS activity and the number of capillaries per
fibre. These changes were associated with a lower blood
lactate and RER during moderate and intense submaximal
running reflecting a greater fat oxidation, which are typical
observations after a period of endurance training (Casaburi
et al. 1987; Beneke and Hutler 2005). Interestingly, soccer
training also resulted in a marked reduction in the oxygen
uptake slow component during intense submaximal

treadmill running as well as an attenuated muscle lactate

and H? accumulation during exhaustive exercise, which
are in line with recent studies investigating the effects of
intense interval training over 8 weeks (Juel et al. 2004;
Krustrup 2004; Mohr et al. 2007). In addition, the soccer
training led to higher lean body mass, quadriceps muscle
mass and mean muscle fibre area, as well as increased
hamstring muscle strength, neither of which were found as
a result of running training. Furthermore, the relative
number of FTx fibres was reduced, as it is typically
observed after a period of strength (Andersen and Aagaard
2000; Kraemer et al. 1995) or endurance training (Coggan
et al. 1992; Kraemer et al. 1995). Thus, soccer training
resulted in superior physiological adaptations when compared with running training.
The differences in the performance development and
adaptations with the two types of training are probably
related to differences in the activity profile, fibre type
recruitment and metabolic response during training. The
running was continuous with the same running speed
during the 1 h of training. In contrast, the recreational
soccer was intermittent with activity changes every 4 s and
included multiple intense actions consisting of *100
intense runs, *60 backwards/sideways runs, 11 tackles, 16

123

1256

Eur J Appl Physiol (2010) 108:12471258

Rest
Exhaustion
3 min recovery

A 13

*
*
6.5 km/h

Blood lactate (mM)

*
8.0 km/h

9.5 km/h

+
SO

11.0 km/h

Incremental test
peak value

0 wks
4 wks
12 wks

*+

*
RU

0
6.5 km/h

8.0 km/h

9.5 km/h

11.0 km/h

Incremental test
peak value

$
$

60

40

20

SO
12 wks

RU
0 wks

RU
12 wks
Rest
Exhaustion
3 min recovery

100

80

60

40
$

SO
0 wks

SO
12 wks

RU
0 wks

RU
12 wks

20

0 wks
4 wks
12 wks

Rest
Exhaustion
3 min recovery

7.4
7.3

4
+

Blood lactate (mM)

13
12
11
10

CO
0
6.5 km/h

8.0 km/h

9.5 km/h

11.0 km/h

Incremental test
peak value

Fig. 5 Blood lactate during treadmill walking at 6.5 km/h and

treadmill running at 8.0, 9.5 and 11 km/h performed before as well as
after 4 and 12 weeks of soccer practice (a; SO, n = 12) and running
(b; RU, n = 10) for untrained men, or continuation of an inactive lifestyle (c; CO = 10). Mean SEM are presented. * Significant
difference from 0 weeks. Significant difference from 4 weeks.
?
Significant difference from CO

shots, 15 dribbles and 67 turns interspersed by low intensity recovery periods. Previous studies from our laboratory
have shown that fast-twitch fibres are heavily recruited
during such activities in contrast to moderate intensity

123

$
80

SO
0 wks

13
12
11
10

100

Muscle CP (mmol/kg d.w.)

*+

Muscle lactate (mmol/kg d.w.)

0 wks
4 wks
12 wks

Muscle pH (-log H )

Blood lactate (mM)

12
11
10

7.2
7.1

6.9

SO
12 wks

RU
0 wks

7.0

6.8
6.7
0.0
SO
0 wks

RU
12 wks

Fig. 6 Muscle lactate (a), CP (b) and pH (c) before and immediately
after the Yo-Yo intermittent endurance level 2 test (Yo-Yo IE2) and
3 min into the recovery period before and after 12 weeks of soccer
practice (SO, n = 12) and running (RU, n = 10) for untrained men.
Mean SEM are presented. $Significant difference from before the
test. Significant difference from exhaustion. * Significant difference
from before the training period

Eur J Appl Physiol (2010) 108:12471258

continuous exercise (Krustrup 2004; Krustrup et al. 2004,

2006b) and this may partly explain the differentiated
adaptations in intermittent exercise and sprint performance
as well as muscle hypertrophy and FTx fibre transformation. The average heart rate was the same during the two
types of training. However, the heart rate was above 90%
of individual HRmax for 20% of the time during the soccer
training when compared with 1% in the running group.
Muscle and blood lactate values were also higher during
soccer training than during running, reflecting a larger
anaerobic energy turnover. These findings indicate that the
high exercise intensity during soccer training, rather than
the duration, is important in order to obtain performance
enhancement after the initial 4 weeks of the training period. In accordance, Jensen et al. (2009) observed that one
additional 30-min aerobic high intensity training session a
week consisting of small-sided games improved the intermittent exercise performance of experienced players during
the season. Furthermore, a number of studies have shown that
endurance athletes can improve their performance by
including activities with intensities higher than corresponding
to VO2max (Esfarjani and Laursen 2007; Iaia et al. 2008).
In summary, soccer training appears, in contrast to
continuous running, to be effective in stimulating to performance development throughout a 12-week training
period and to cause significant cardiovascular and muscle
adaptations including muscle growth and elevated muscular strength. This may be related to the finding of many
intense actions and a significant stimulation of the anaerobic system during the small-sided games. Thus, recreational soccer appears to be an effective type of training
leading to significant performance improvements and
muscle adaptations.
Acknowledgments We thank all the participants and Rikke Leihof,
Thomas Bonne, Mihai Ursta Relu, Rasmus Bischoff, Mads Bendiksen
and Thomas Gunnarsson for excellent technical assistance. The study
was supported by the Danish Ministry of Culture (Kulturministeriets
Udvalg for Idrtsforskning) and by the Danish Football Association
(Dansk Boldspil-Union).

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