Journal of Sports Sciences, October 2005; 23(10): 1101 1109

Changes in muscle activity with increasing running speed

LA
INEN, JANNE AVELA, & PAAVO V. KOMI
HEIKKI KYRO
Neuromuscular Research Centre, Department of Biology of Physical Activity, University of Jyvaskyla, Jyvaskyla, Finland
(Accepted 9 October 2004)

Abstract
Electromyographic (EMG) activity of the leg muscles and the ground reaction forces were recorded in 17 elite male middledistance runners, who performed isometric maximal voluntary contractions (MVC) as well as running at different speeds.
Electromyograms were recorded from the gluteus maximus, vastus lateralis, biceps femoris, gastrocnemius and tibialis
anterior. The results indicated that the averaged EMG (aEMG) activities of all the muscles studied increased (P 5 0.05) with
increasing running speed, especially in the pre-contact and braking phases. At higher speeds, the aEMG activities of the
gastrocnemius, vastus lateralis, biceps femoris and gluteus maximus exceeded 100% MVC in these same phases. These
results suggest that maximal voluntary contractions cannot be used as an indicator of the full activation potential of human
skeletal muscle. Furthermore, the present results suggest that increased pre-contact EMG potentiates the functional role of
stretch reexes, which subsequently increases tendomuscular stiffness and enhances force production in the braking and/or
propulsive phases in running. Furthermore, a more powerful force production in the optimal direction for increasing running
speed effectively requires increased EMG activity of the two-joint muscles (biceps femoris, rectus femoris and
gastrocnemius) during the entire running cycle.

Keywords: Electromyography, ground reaction force, stretch reex

Introduction
Knowledge of the muscle activity patterns of the leg
muscles is important for understanding the kinematics and kinetics of running. Highly developed
motor control with proper timing of movements is
required for effective force production. Thus, muscle
activity patterns should primarily be examined with
respect to their function during ground contact. This
contact phase represents the stretch shortening
cycle action for the leg extensor muscles, which is
benecial for force output (Cavagna, 1977; Komi,
2000).
However, increased force production with increasing running speed appears mainly to be due to
increased pre-activity of the leg extensors (Komi,
Gollhofer, Schmidtbleicher, & Frick, 1987) and the
earlier onset of extensor activity (Nilsson, Thorstensson, & Halbertsma, 1985). The activation of the
leg extensor muscles before foot contact appears to
be pre-programmed (Dietz, Schmidtbleicher, &
Noth, 1979; Melvill-Jones & Watt, 1971), where
amplitude and timing depend on foot position and
the expectancy of the moment during ground contact

(Dietz, 1992) and on running speed (Komi et al.,

1987). Some of the programmed electromyographic
(EMG) patterns may be inborn and generated on a
spinal and/or a brainstem level (Dietz, 1992).
The selection of an appropriate pre-programmed
pattern by the supraspinal centres depends on
information delivered via afferents from muscles,
joints and associated tissues (Dietz, 1992). The
group I afferent reex system has a very important
role in controlling posture and locomotion, because
its gain can be modulated by pre-synaptic inhibition
and by changes in muscle spindle sensitivity through
the fusimotor system (Dietz, 1992; Loeb & Hoffer,
1985; Stein, 1978). Stretch reexes regulate tendomuscular stiffness and increase force output (Hoffer
& Andreassen, 1981). In line with this notion, stretch
reexes have been shown to potentiate dramatically
the force production after an electromechanical delay
of 13 15 ms (Nicol & Komi, 1998).
During the contact phase in running, EMG
amplitude of the leg extensor muscles increases with
increasing running speed (Kyrolainen, Komi, &
Belli, 1999; Mero & Komi, 1987). More specically,
the EMG activity of the hip extensors increases

Correspondence: H. Kyrolainen, Department of Biology of Physical Activity, University of Jyvaskyla, FIN-40100 Jyvaskyla, Finland. E-mail: heikki@sport.jyu.
ISSN 0264-0414 print/ISSN 1466-447X online 2005 Taylor & Francis
DOI: 10.1080/02640410400021575

1102

H. Kyrolainen et al.

markedly (Kyrolainen et al., 1999; Simonsen,

Thomsen, & Klausen, 1985), and the onset of
extensor muscle activity begins earlier (Nilsson et
al., 1985) that is, the duration of pre-activity
increases. Furthermore, the EMG activity of the
gastrocnemius and soleus muscles has been shown to
be higher in running than in maximal isometric tests
(Mero, Kuitunen, & Komi, 2001). The present study
was designed to examine the EMG activity of a
number of leg muscles at several submaximal
running speeds in relation to the respective values
recorded either during maximal running or isometric
maximal voluntary contractions (MVC), as these
conditions should provide different possibilities for
reex-induced activation.

Methods
Participants
Seventeen elite male (age 23 + 3 years, height
1.80 + 0.05 m, body mass 66.9 + 6.0 kg, body fat
7.4 + 1.9%) middle-distance runners volunteered
as participants for the present study. All runners
were fully informed of the procedures and possible
risks of the experiment, and they provided written
consent to participate in it. The study received
approval from the Ethics Committee of the
University of Jyvaskyla.
Procedure
After a 30-min warm-up, the participants were asked
to perform two maximal voluntary contractions in
different conditions separately for three muscle
groups: knee extensors, plantar exors and knee
exors. Before the tests, the participants practised
the isometric force production three to four times.
During the tests, they were verbally encouraged to do
their best.
After 10 min recovery, the participants ran ve
submaximal running bouts on an indoor track
(Jyvaskyla, Finland) followed by three maximal
sprints. The submaximal tests began with 3 min
running at a predetermined constant speed of
4.0 m s71 (48 + 3% of the maximal speed), followed by speeds of 5.0 m s71 (58 + 4%) and
5.5 m s71 (66 + 3%) with 8 min recovery between
each 3-min bout. After 8 min recovery at the end of
these three bouts, they ran 1 min at each of two
constant speeds of 6.0 m s71 (73 + 4%) and
7.00 m s71 (82 + 4%), with again 8 min recovery
between bouts. Running was performed on a 200-m
track so that the runners followed the light system so
as to maintain a constant speed (Proton, Naakka
Inc., Finland). Finally, after 15 min recovery, they
ran three maximal sprints over a distance of 30 m,

with a run-up distance that was individually selected.

The mean ( + s) maximal speed of the participants
was 8.50 + 0.57 m s71.
Measurements
In the MVC test for the knee extensors, force was
measured at a standardized knee angle of 1078.
Electromyographic activity (ME3000P, MEGA,
Kuopio, Finland) was simultaneously recorded telemetrically from the vastus lateralis and the rectus
femoris using bipolar surface electrodes (Beckman
miniature skin electrodes, 650437, California, USA).
The EMG activity of the tibialis anterior was
recorded only during running. The electrodes, which
had a centre-to-centre distance of 20 mm, were
placed longitudinally over the muscle bellies between
the centre of the innervation zone and the distal
tendon of each muscle. The EMG signal amplication was 500 (Glonner Biomes 2000; cut-off
frequency 360 Hz/3 dB71), and it was digitized
simultaneously with the force records at a sampling
frequency of 1 kHz. The maximal EMG activities of
the other muscles studies were measured in maximal
isometric conditions without force recordings. The
EMG activity of the biceps femoris was measured in
the prone position with a knee angle of 908 and a hip
angle of 1808. The EMG activity of the gastrocnemius was recorded during a maximal plantar exion
with the knee fully extended (1808) and an ankle
angle of 908. The voluntary EMG activity of the
gluteus maximus was measured with the hip joint
extended isometrically and the the knee fully
extended.
In all running tests, the average speed was
measured and controlled by photocells (Newtest,
Oulu, Finland). During each lap, ground reaction
forces were measured by a 10-m long force platform
(TR-testi, Finland and Kistler, Switzerland: natural
frequency 5 150 Hz, linearity 5 1%, crosstalk
4 2%). The EMG activity of the muscles was
recorded as in the isometric force tests.
MVC and running biomechanics
In the MVC test for the knee extensors, maximal
force and averaged EMG (aEMG) of the vastus
lateralis, together with the entire force time curve,
were analysed. Maximal aEMG activities of the other
muscles studied were determined for a period of 1 s.
Individual ground reaction forces and aEMG activities from 5 10 contact phases at each running
speed were selected for further analysis. To synchronize these signals, a vertical force signal of 50 N was
used to identify and trigger the beginning and the
end of the contact. Ground reaction forces were
divided into braking and propulsive phases according

Muscle activity in running

to the orientation of the horizontal force (Mero &
Komi, 1987). The pre-activity was analysed 100 ms
before ground contact (Komi et al., 1987; Kyrolainen, Belli, & Komi, 2001). All aEMG activities were
compared with the respective aEMG activities
measured both in the MVC and maximal running
conditions (Figure 1).
Statistical analysis
Owing to high inter-individual variation, especially
for the EMG variables, non-parametric statistical
techniques were adopted in the present study.
Friedmans two-way analysis of variance (chisquare) was used to test the effects of experimental
conditions (MVCs and running speeds), and the
Wilcoxon matched-pairs signed-rank test was used
for the repeated measures. In addition, Spearmans
rank order correlation coefcient was used to
determine the relationship between the measured
variables. All data are presented as the mean + standard deviation.

Figure 1. An example of the activity recorded during isometric

maximal voluntary contractions (MVC) and during running. The
arrows indicate the beginning of force production.

1103

Results
Neuromuscular capacity
The mean maximal force of the knee extensors was
1358 + 312 N, the rate of force development was
5653 + 2046 N s71, and the maximal aEMG activity of the vastus lateralis was 473 + 185 mV.
Compared with maximal voluntary contractions,
the aEMG activities during maximal running were
higher (P 5 0.001) in the pre-contact (753 +
253 mV) and braking phases (618 + 232 mV), but
clearly lower (P 5 0.001) in the propulsive phase
(155 + 73 mV). Table I presents the aEMG activities
of all muscles.

Figure 2. Vertical (Fz) and horizontal (Fy) ground reaction forces

at different running speeds. The bold lines represent maximal
running.

Table I. EMG activities (mV) measured during maximal voluntary contractions (MVC) and in the different phases of maximal running
Running phases

Gastrocnemius
Vastus lateralis
Biceps femoris
Rectus femoris
Gluteus maximus
Tibialis anterior

MVC

Pre-activity

Braking

Propulsive

459 + 251
473 + 185
398 + 107
363 + 152
215 + 83

802 + 272***
753 + 253***
343 + 127
269 + 174
362 + 188***
266 + 108

868 + 280***
618 + 232*
442 + 123
258 + 154
415 + 235***
163 + 62

590 + 214***
155 + 73***
116 + 37***
79 + 59***
147 + 111
132 + 58

Note: Statistically signicant differences in EMG between the running phases and MVC: *** P 5 0.001, * P 5 0.05.

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H. Kyrolainen et al.

Running kinematics and kinetics

The contact times shortened gradually from
0.203 + 0.011 s (braking phase of 0.098 + 0.008 s)
at a speed of 4.0 m s71 to 0.112 + 0.007 s (braking
phase of 0.053 + 0.005 s) at maximal speed
(w2 = 85.00, P 5 0.001). At the same time, step
frequency increased from 2.82 + 0.13 to 4.16 +
0.26 Hz (w2 = 85.00, P 5 0.001) and step length
from 1.51 + 0.10 to 2.12 + 0.15 m (w2 = 81.35,
P 5 0.001).
Figure 2 shows that the magnitude of the ground
reaction forces and the rate of force production
increased with increasing running speed. The maximal vertical force was 2012 + 142 N in the braking

phase at a speed of 4.0 m s71, which increased to

2366 + 364 N (w2 = 63.77, P 5 0.001) at maximal
speed. Simultaneously, the maximal horizontal force
increased in the same phase from 339 + 33 to
830 + 210 N (w2 = 77.74, P 5 0.001). The average
resultant forces, which were related to body weight,
increased from 2.1 + 0.2 to 2.5 + 0.2 (w2 = 73.20,
P 5 0.001) in the braking phase of ground contact.
However, only minor changes in the direction of force
(83.2 + 0.88 vs. 80.9 + 1.48 from the horizontal)
were observed. In the propulsive phase, the relative
resultant force increased from 1.5 + 0.1 to 1.8 + 0.1
(w2 = 61.13, P 5 0.001), and its direction inclined
slightly more towards the horizontal (81.8 + 0.78 vs.
76.9 + 1.38) (w2 = 69.91, P 5 0.001).

Figure 3. (Top) Muscle activity patterns of the vastus lateralis at a speed of 4.0 m s71 (thin line) and at maximum speed (bold line).
(Bottom) Mean ( + s) relative EMG activities for three functional phases. The dashed horizontal lines indicate the maximum (100%) EMG
measured during maximal voluntary contractions (MVC), and the dashed vertical line represents the beginning of the ground contact. Chisquare indicates statistically signicant changes with increasing running speed.

Muscle activity in running

Running EMGs
The gluteus maximus was most active in the late
swing and braking phases of contact. Its aEMG
activity increased (P 5 0.001) during the pre-contact
(174 + 108 mV at a running speed of 4 m s71 vs.
362 + 188 mV at maximal speed) and braking phases
(150 + 83 mV vs. 415 + 235 mV) with increasing
running speed. Figure 3 demonstrates that the aEMG
activity of the vastus lateralis increased (P 5 0.001 to
0.01) in all phases of running, with the greatest
increases in the pre-contact and braking phases. In
these phases, the activity of the vastus lateralis at the
higher running speeds exceeded that of the maximal
voluntary contractions, while in the propulsive phase
aEMG activity was below 50% MVC.

1105

The muscle activity patterns of the two-joint

muscles, the rectus femoris and biceps femoris,
demonstrate clear changes due to running speed
(Figures 4 and 5). The aEMG activity of these
muscles increased (P 5 0.001 to 0.05) both in the
swing and contact phases with increasing running
speed. The mean relative aEMG activity did not
reach the level seen during maximal voluntary
contractions for the rectus femoris. For the biceps
femoris, however, aEMG activity in the braking phase
reached at maximal speed the same level as that
measured during maximal voluntary contractions.
The aEMG activity of the gastrocnemius increased
in both the pre-contact and braking phases
(P 5 0.001). Figure 6 shows that at speeds above
6.0 m s71, aEMG activity in the pre contact phase of

Figure 4. (Top) Muscle activity patterns of the rectus femoris at a speed of 4.0 m s71 (thin line) and at maximum speed (bold line).
(Botom) Mean ( + s) relative EMG activities for three functional phases. The dashed horizontal lines indicate the maximum (100%) EMG
measured during maximal voluntary contractions (MVC), and the dashed vertical line represents the beginning of the ground contact. Chisquare indicates statistically signicant changes with increasing running speed.

1106

H. Kyrolainen et al.

Figure 5. (Top) Muscle activity patterns of the biceps femoris at a speed of 4.0 m s71 (thin line) and at maximum speed (bold line).
(Botom) Mean ( + s) relative EMG activities for three functional phases. The dashed horizontal lines indicate the maximum (100%) EMG
measured during maximal voluntary contractions (MVC), and the dashed vertical line represents the beginning of the ground contact. Chisquare indicates statistically signicant changes with increasing running speed.

running was greater than that seen during maximal

voluntary contractions. In the braking phase, the
aEMG activity of the gastrocnemius was higher
already at the slowest speed than during maximal
voluntary contractions. In the propulsive phase, the
relative EMG activity of the gastrocnemius was higher
at maximal speed than during maximal voluntary
contractions. The aEMG activity of the tibialis
anterior increased (P 5 0.001) in the middle of the
swing phase and at the beginning of the contact phase.

muscles during the braking phase of running. Their

activity is in many cases well above that recorded
during maximal voluntary contractions, suggesting
that such contractions cannot be used as an indicator
of the full activation potential of human skeletal
muscle. Thus, features of central programming and
reex regulation seem to differ between isometric
maximal voluntary contractions and stretch shortening cycle exercises such as running.
Role of pre-activity and the proprioceptive reex system

Discussion
The results of this study emphasize the importance of
pre-activity and the activity of the leg extensor

The pre-activity increased in all muscles studied with

increasing running speed, and the relative EMG
amplitudes of the vastus lateralis and gastrocnemius

Muscle activity in running

1107

Figure 6. (Top) Muscle activity patterns of the gastrocnemius at a speed of 4.0 m s71 (thin line) and at maximum speed (bold line).
(Botom) Mean ( + s) relative EMG activities for three functional phases. The dashed horizontal lines indicate the maximum (100%) EMG
measured during maximal voluntary contractions (MVC), and the dashed vertical line represents the beginning of the ground contact. Chisquare indicates statistically signicant changes with increasing running speed.

were higher at the faster speeds than observed during

maximal voluntary contractions. For the gastrocnemius it has been demonstrated that the higher the
running speed, the higher the pre-activity (Komi et
al., 1987). In the present study, the higher preactivity was related to higher activity in the subsequent braking phase of the contact (Figure 7). This
relationship became stronger as running speed
increased. Therefore, pre-activity appears to be a
preparatory requirement both for the enhancement
of EMG activity during the braking phase and for the
timing of muscular actions with respect to ground
contact.
Pre-activity is assumed to increase the sensitivity of
the muscle spindle through enhanced alpha gamma
coactivation, leading to the potentiation of stretch

reexes (Gottlieb, Agarwal, & Jaeger, 1981). In other

words, during the braking phase of running, changes
in muscle spindle sensitivity increase activity of the
type Ia afferents and subsequently facilitate the amotoneurones resulting in increased EMG activities
of the knee extensor and plantar exor muscles with
increasing running speed. As a consequence, centrally programmed pre-landing activity (Melvill-Jones
& Watt, 1971) seems to enhance the stiffness of the
tendomuscular system to tolerate and absorb high
impact loads at the beginning of ground contact (e.g.
Gollhofer & Kyrolainen, 1991; Hoffer & Andreassen,
1981). Stretch reexes can enhance force production,
alter the latency time and electromechanical delay, as
Nicol and Komi (1998) have demonstrated under
laboratory conditions, not only in the braking phase

1108

H. Kyrolainen et al.
(Fellows et al., 1993). Utilizing partial blockade of
type Ia afferents by ischaemia, Dietz et al. (1979)
suggested that spinal stretch reexes, which play an
important role in stiffness regulation (Hoffer &
Andreassen, 1981), could affect powerful force
production in the propulsive phase. The increased
muscle activity of the leg extensors in the braking
phase of the contact is also a prerequisite for efcient
storage of elastic energy (Komi & Gollhofer, 1997).
When running at slower speeds (e.g. during the
marathon), the reex-induced force enhancement
will already have a functional effect during the
braking phase, while at maximal speeds its net reex
contribution will occur at the end of the braking
phase or even in the propulsive phase (Komi, 2000).
As a consequence of the high pre-landing and
braking activity of the leg extensor muscles, unnecessary yielding of the runner during the braking
phase might be prevented.
Role of the biarticular muscles

Figure 7. Relationships between relative EMG activity in the precontact and braking phases of the vastus lateralis (top) and
gastrocnemius (bottom) muscles at a speed of 4.0 m s71 and at
maximum speed.

of running but also in the propulsive phase depending

on the duration of contact in running. It is thought
that the central nervous system is capable of full
activation of all motor units during maximal isometric exercise (e.g. Rice, Andrews, & BiglandRitchie, 1991). However, based on the present results
and on those of earlier studies (Fellows, Domges,
Topper, Thilmann, & Noth, 1993; Mero et al., 2001),
this may not be the case.
Powerful running requires a combination of
inherent muscle/joint stiffness and enhancement of
muscle force resulting from segmental stretch
reexes (Dietz et al., 1979; Kuitunen, Komi, &
Kyrolainen, 2002). Stretching the tendomuscular
system changes muscle spindle sensitivity (Matthews
& Stein, 1969) causing increased type Ia afferent
activity (Matthews, 1984), and subsequently facilitation of motor output (force enhancement) due to
increased EMG activity (Houk, 1979). These basic
ndings are in line with the results of the running
study of Dietz et al. (1979), who observed that the
electrical activity of the gastrocnemius increased
sharply about 40 ms after ground contact in running.
This short latency reex component is mediated by
afferent inow via type Ia muscle spindle afferents

The biarticular muscles are suggested to contribute

to the transfer of power from the proximal to the
distal joints (e.g. Bobbert & Van Ingen Schenau,
1988; Jacobs, Bobbert, & Van Ingen Schenau, 1996)
and nally to the ground. In the present study, the
EMG activity of the rectus femoris increased
markedly with increasing running speed in two
phases: rst, just before and at the beginning of
contact when its contribution as a knee extensor
increased; and second, at the beginning of the swing
phase when it started to powerfully ex the hip joint.
For the biceps femoris, the EMG activity increased
both in the latter part of the swing phase and in the
contact phase. Simonsen et al. (1985) have interpreted this phenomenon as the hamstring muscles
releasing elastic energy during ground contact, which
is stored during the ight phase when they function
eccentrically. Thus, the biceps femoris contributes to
more powerful extension of the hip joint. Consequently, in the subsequent propulsive phase, the
hamstrings drive the body more powerfully in the
forward direction when the propulsive force increases, resulting in a more inclined resultant force
towards the horizontal.
In general, the biarticular muscles perform eccentric work during the swing phase and concentric
work during the whole ground contact phase.
Prilutsky, Gregor and Ryan (1998) have found a
close relationship between EMG activity and joint
moments during the swing phase (without an
external force applied to the foot) of walking and
running. They suggested that the observed coordination of the rectus femoris and hamstring muscles can
be regulated without a stance-specic modulation of
cutaneous afferent input from the foot.

Muscle activity in running

In conclusion, isometric maximal voluntary contractions may not be a good indicator of the full
activation potential of human skeletal muscle. The
EMG activity of the leg extensor muscles during
running exceeds considerably that recorded during
isometric maximal voluntary contractions. In running, increased pre-contact EMG activity is essential
in particular for higher reex activity and, therefore,
for better stiffness regulation of the muscle. Furthermore, a more powerful force production with
increasing running speed requires drastically increased EMG activity of the two-joint muscles
during the entire running cycle.
Acknowledgements
The authors would like to express their gratitude to
Jarkko Finni, Markku Ruuskanen, Juha Kuosa, Sirpa
Roivas, Marja-Liisa Romppanen and Pirkko Puttonen for their help with data collection and Esko
Levalahti for his assistance in the statistical analysis.
This study was supported by grants #170/722/95,
#104/722/96 and #96/722/97 from the Ministry of
Education, Finland.

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