Acta Physiol Scand 1981, 111: 135-140

Prestretch potentiation of human skeletal muscle

during ballistic movement
CARMELO BOSCO, PAAVO V. KOMI and AKIRA I T 0
Department of Biology of Physical Activity, University of Jyvaskyla, SF-40 100 Jyvaskyla 10, Finland

BOSCO, C., KOMI, P. V. & ITO, A.: Prestretch potentiation of human skeletal muscle
during ballistic movement. Acta Physiol Scand 1981, 111: 135-140. Received 14 April 1980.
ISSN 0001-6772. Department of Biology of Physical Activity, University of Jyvaskyla,
Finland.

The conditions associated prior to and during the transition from prestretch to shortering
may have considerable influence on the final performance of muscle. In the present study
male subjects of good physical condition performed vertical jumps on the force-platform
with and without preliminary counter movement. In the counter movementjump (CMJ) the
amplitude of the knee bending, velocity of the prestretch and the force attained at end of
prestretch were the primary parameters of interest. In addition the coupling time indicating
the transition from the eccentric (prestretch) phase to the concentric phase was recorded
from the angular displacement and reaction force curves. In the final calculation the
mechanical performance parameters of CMJ were always compared with those of the jumps
performed without counter movement. The results indicated in general first that CMJ
enhanced the average concentric force and average mechanical power by 423 N (66%) and
1158W (81 %), respectively. This potentiation effect was the higher the higher was the force
at end of prestretch (p<O.001). Similarly, the prestretch speed (p<O.OOl) and short coupling
time (pcO.01) were associated with enhanced performance during the concentric phase.
The average coupling time was 23 ms. The results are interpreted through changes in the
prestretch conditions to modify the acto-myosin cross-bridge formation so that the storage
and utilization of elastic energy is associated with high prestretch speed, high eccentric
force and short coupling time. The role of the reflex potentiation is also suggested as
additional enhancement of the final performance.

Key words: Muscle mechanics, stretch-shortening cycle, muscle elasticity, prestretch

potentiation.

Stretching of an activated muscle prior to its short-
ening enhances its performance during the concen-
tric phase. This phenomenon, which has been ob-
served both in animal (Cavagna et al. 1965, 1968)
and in human experiments (e.g. Cavagnaet al. 1968,
Cavagna et al. 1971, Thys et al. 1972, Asmussen &
Bonde-Petersen 1974a, Komi & Bosco 1978, Bosco
& Komi 1979) has been interpreted to be primarily
due to the release of elastic energy stored in the
series elastic elements of the muscle during stretch.
The capacity of the muscle’s ability to store and
utilize elastic energy may depend on the speed of
prestretch, on the final muscle length and on the
force developed at end of prestretch (Cavagna et al.
1%8). On the other hand, movement amplitude per
se has been demonstrated to influence the perfor-
mance characteristics so that high mechanical effi-
ciency was associated with small angular displace-
ment during knee bending exercises (Thys et al.
1975, Asmussen & Bonde-Petersen 1974b). Similar-
ly the small movement amplitude has been dem-
onstrated to enhance the force and power output
of the skeletal muscles in a test where the subjects
performed vertical jumps on the force-platform with
and without counter movement (Bosco & Komi
1980). Utilizing these similar force-platform
techniques, the present study was undertaken to
examine further the influence of prestretch speed,
prestretch amplitude, the force attained at end of
prestretch and the transition (coupling) time be-
tween the eccentric and concentric phases on the
final performance. It was thought that these factors
would also be connected with the modifications of
the cross-bridge formation (Huxley & Simmons

Acta Phvsiol Scand 111

I36 Carmelo Bosco et al.

A IMPACT TAKE OFF Table 1. Meun (kS.D.1 wsults of the mechanical

wricibles stiidied

Parameters Mean k S.D.

Knee angle amplitude, a (degrees) 71.47f21.36

Prestretch range, aps(degrees) 18.84k1.52
Prestretch speed, P,, (rad/sec) 4.39-1 1.87
B Prestretch time, T,, (ms) 85.03f35.52
Coupling time. T, (ms) 22.97+ 14.67

I I 5 Rad
Force at end of stretch.
-F, (N)
Average prestretch force,
-F (N)
2 484.64+901.87

1 197.00i470.92
Potgntiation of average force,
hF (N) 423.15k236.85
Potentiation of average power,
AW (Watts) 1 158.08k546.09

0 positive work phase @. The force-time curve and the

COUPLING
TIME
0 0.5 1.0 i s 1
I ,
F.i.y. I . Examples of the vertical force-time curve and an-
gular displacement during static jump (SJ) and counter
movement jump (CMJ). In SJ the force-time curve during
the take off reads positive values as indicated by $. In
CMJ the vertical ground reaction force is first negative
during the unweighing phase (UW) but reads positive
values both during the decelerating phase (8)and posi-
tive work phase (63). The coupling time in the angular
displacement record indicates the transition period be-
tween the eccentric and concentric phases.
1971) and consequently influence the final perfor-
mance in the concentric phase.
flight time (taJ gave the basis for calculation of the fol-
lowing parameters: vertical velocity at take-off -(Vv),
average force (F) and average mechanical power (W). In
CMJ condition both the average force during the de-
celerating phase (-F) and the instantaneous force de-
veloped at the end of the stretch (-F,) were computed.
The vertical velocity at take-off was obtained from the
formula:
v, = 112 t,,,xg (1)
in which g=acceleration of gravity (9.81 m/s2).
Average force (F) was then computed with the following
formuta:
E= F=m x V,
t
in which [=the respective time of the force-time curve,
m=mass of the subject, V,=final vertical velocity at
take-off.
Total mechanical work (W) was calculated according to
the mechanical energy changes during contact as follows

W = 1/2 mV2+mgh (3)

METHODS
Subjects. 14 male well trained power athletes were used as
subjects (age 22.93rt3.54 years. height 181.64-18.21 cm
and weight 74.14i7.03 kg). All the subjects tested were
familiar with the test procedure used. This consisted of
performing maximal vertical jump on the force-platform
(Komi et al. 1974) with and without counter movement.
These performances are called respectively counter
movement jump (CMJ) and squatting jump (SJ).
Each jump produced a vertical force-time curve,
examples of which are given in Fig. 1 A-C. From SJ condi-
tion the force recorded during the contact is always
positive, whereas in CMJ it is first negative (below the
body weight) during the unweighing phase (UW) but
stays positive both during the deceleration phase 8 and
where m=mass of the subject, v=final velocity at take-
off, h=rise of the center of gravity (C.G.) from the lowest
point during contact, and g=acceleration of gravity (9.81
m/sz).
The final velocity and rise of C.G. were computed by
integration of the acceleration from the force-time curve.
The final vertical velocity was similar (rtO.01 m/s) to the
V,. calcu!ated from formula (1). The average mechanical
power w) was given by formula:
*=w t (4)
In order to examine the movement amplitude during SJ
and CMJ, an electrogoniometer (Elgon) was attached to
the lateral side of the subject knee joint. Elgon recorded

.-t<tO P / i Y \ l , J l .S"Itld Ill

 Prestretch potentiation of muscle force 137

Table 2 . Correlation matrix for the various parameters calculated from the force-time curve and Elgon
records of 59 observations
1 2 3 4 5 6 7 8

Knee angle amplitude (a) 1

Prestretch range (aDs) 2 39
Prestretch speed (PsJ 3 51 17
Prestretch time (TPJ 4 75 50 -68
Coupling time (T,) 5 46 29 -21 52
Force at end of stretch (-FJ- 6 -67 -35 61 -14 -41
Average prestretch force (-F) - 7 -53 -47 49 -16 -47 76
Potentiation of average force (FL 8 -61 -13 53 -62 -35 51 43
Potentiation of average power (W) 9 -47 -23 33 -54 -41 48 37 81

the angular displacement, which ranged from 27” to 105”in
all subjects. From this total angular displacement (a)
which included both the unweighing and prestretch phases
the latter could also be computed as denoted by apB in Fig.
1. This phase is a position of the decelerating phase 0 of
the force-time curve and here the muscles are actively
resisting the imposed stretch and the phenomenon is
therefore called as prestretch. The shaded area of the Fig.
1 represents the position of the Elgon record where the
knee angle stays the same and supposedly no change
occurs in the knee extension muscles. This phase is called
coupling time between the prestretch (eccentric) and short-
ening (concentric) phases. The average prestretch speed
was then obtained by dividing aPsby the respective time
(tPS).
RESULTS
In presenting the results the comparison is made in
performance difference between CMJ and SJ condi-
tions, which utilized similar movement ranges
around the knee joint in the concentric phase. This
was possible because CMJ was performed always
first and the starting angle for SJ was obtained from
the Elgon record of CMJ.
Table 1 presents first the absolute mean (+S.D.)
values for the selected mechanical attributes. As is
seen from this table the average prestretching time
in CMJ was 85.03k35.52 ms, and the coupling time
23.0k14.7 ms. During this phase the force values
were 2084.6k901.9 N. As compared to SJ the
potentiation for the average concentric force (AF)
and average concentric power (AW) were
423.2k236.9 N and 1 158.1k546.1 W, respectively.
These correspond to relative changes of 66 and
81 %.
Correlation analysis revealed that several
mechanical parameters were correlated with the
“potentiation” variables of A F and AW (Table 2 ) .
For example the value of F was correlated posi-
tively by prestretching speed ( r=0.53,P <O. 00 1 ;Fig.
2 ) , negatively by coupling time (r=-0.35, P<O.Ol),
and positively by instantaneous force at end of
stretch (r=0.51,P<O.OOl). Similarly Fig. 3 presents
how change in average power output (AW) was
correlated by the respective variables.
The coupling time, which denoted the “isometric”
transition time from the eccentric phase was the
longer the higher was the movement amplitude
(r=0.46, P<O.OOl; Fig. 4 ) . Correspondingly its
duration was influenced by the force attained at the
end of stretch ( r = -0.47, P<O.OOl; Fig. 5 ) .
DISCUSSION
The present results indicated clearly, and in agree-
ment with previous studies (e.g. Cavagna et al.
1971, Asmussen & Bonde-Petersen 1 9 7 4 ~Komi ~
& Bosco 1978) that utilization of the stretch-short-
ening cycle enhances the performance of muscle
over that of the pure concentric contraction. This
was observed when the performance variables of
CMJ and SJ were cornpxed. Mean increases of
66% and 81 % in average force and power output,
respectively, are substantial and their absolute val-
ues were highly dependent on the speed during the
decelerating phase. This finding must be related to
the concept of short range stiffness (Rack &West-
bury 1974) and consequently it should be related to
the duration of the “coupling time” between the
eccentric and concentric phases. In addition, fast
stretch associated with small amplitude is likely to
cause the final eccentric force be high (Joyce et al.
1974) and therefore due to increased stiffness make
the transition from the eccentric to concentric
phase take place more quickly.
In the present result this hypothesis is supported
by the negative correlation (P<O.OOl) between the
force at end of stretch and the coupling time. This

Acta Physiol Scand I l l

138 Carmelo Bosco et al.

PRESTRETCH
IR a d / s 1
.. . In.5O 1
SPEED
y = 2 672 - 0 OOL x
r . 5 3 ( P= 001) .
..
0

..
.
/

-
0
A (,MI aw (watt 1
0 LOO 800 1200 1000 2000

FORCE AT END
L2W - OF PRESTRETCH

3200 -

2200 -

1200 -

1
0 LOO
.,
800
A F IN)
1200

y I

n : 59
320.18 - , 21x

. I

t
0 LOO 800 I200
0
0
. 1000
- A5 ( w a t t )
2000

F;g. 2. Prestretch speed (Pss),instantaneous negative
force ( - F , ) developed at end of prestretch phase, and
coupling time (T,) calculated from CMJ are shown as
function of average force difference between CMJ and SJ
(AF).
Fig. 3. Prestretch speed (PSS),instantaneous negative
force (-F,) developed at end of prestretch phase, and
coupling time (T,) calculated from CMJ are shown as
function of average positive power difference between
CMJ and SJ (AW).

would naturally have consequences in fast ballistic
movements. The short coupling time is likely to
have also other consequences. It has been suggest-
ed that the average duration of the cross-bridge
attachment is 15 ms (Stienen et al. 1978). Now
according to Huxley & Simmons (1971)the heads of
the myosin filaments are rotated backwards during
stretch, against their natural tendency, to the posi-
tion of higher potential energy. These lengthened
cross-bridges can then be detached if the stretch
 Prestretch potentiation of muscle force 139

60 1‘mifl
COUPLING TIME

y
r
i

=
6.0
.16
3.13 x
I Pc.001) b
00
6ol
,;: COUPLING

m a a r = - . 4 1 ( P<OOl)

30 -

-’ 0 0 0 ‘0
0 8.’
.a
.* *. .
.
*a
1 I a
0 20 40 60 80 100 120 a
ANGULAR D I S P L ACE M E NT ( degrees) 0 . a a a

Fig. 4 . Relationship between coupling time (T,) and knee tow 2000 3000 4000 WOO
joint movement amplitude. FORCE DEVELOPED AT END OF PRESTRETCH I N )

Fig. 5. Relationship between coupling time (T,) and in-

position is maintained for too long (Cavagna & Cit-
terio 1974) o r may cause sarcomere “slipping” if the
range of stretch is large (Flitney & Hirst 1978).
Short coupling time will prevent detachment of the
cross-bridges and consequently allow better utiliza-
tion of their stored potential (elastic) energy during
the subsequent shortening phase. This mechanism
would then explain the finding of Cavagna et al.
(1975) that the apparent “potentiation” of the con-
tractile component of the muscle lasts for a limited
amount of time after the end of stretch. Fig. 6 is an
attempt to explain this mechanism schematically.
The above explanation is based on the assumption
that the potentiation effect is primarily of mechani-
cal origin and associated with the elastic behavior
stantaneous negative force (-F,) developed at end of de-
celerating phase during CMJ.
and attachment-detachment cycle of the actin-my-
osin cross-bridges. Other possible explanations are
available. Evidence has been presented that fast
stretch of an active muscle causes substantial
stretch reflex potentiation via Ia afferents from the
muscle spindles (Prochazka et al. 1977). As suggest-
ed by Antoni et al. (1979) this reflex potentiation
with increased motor neuron activity to the contract-
ing muscles would cause the force to be very high
at the end of the eccentric phase. Increased force
would then cause the stiffness of the muscle to
increase. Increased stiffness on the other hand is

SLOW AND

LONG STRETCH

Fig. 6. A schematic model to suggest the attachment and detachment of the acto-my-
osin cross-bridges during the stretch-shorteningcycle associated with different duration
and speed of the prestretch phase. The calculations treat the cross-bridges working in
palallel with each other. This model is based also on the hypothesis of Huxley &
Simmons (1971) and Cavagna & Citterio (1974).

Acta Physiol Scand I I I

140 Carmelo Bosco et al.
likely to make the condition favorable for the short
coupling time within the stretch-shortening cycle. It
is possible that stretching of active muscle causes
reflex potentiation also via cortical loop. There are
indications that this loop may reach the muscle
already within 50 ms from the beginning of stretch
(Iles 1977). This 50 ms corresponds approximately
to the end of the impact (eccentric) phase of the
contact period during fast running. Therefore even
in fast ballistic movements such as sprinting the
contribution of the stretch reflex potentiation is
possible.
A question should be asked what the relative roles
of the two possible mechanics are-elastic and re-
flex-for the final potentiation effect? The present
study was not designed to investigate the reflex
phenomenon. However, it is known from studies
with isolated muscles (Cavagna et al. 1965) that the
pure elastic potentiation is substantial and that in
studies where stretch-shortening cycle has been uti-
lized with intact human muscle (Cavagna et al.
1968. Bosco & Komi 1979) the work-velocity and
force-velocity curves, respectively, shift also to the
right. Therefore it must be concluded that the elas-
tic phenomenon is probably of primary importance
but the role of reflex potentiation cannot be over-
looked especially in fast movements, which utilize
stretch-shortening cycles.
The authors acknowledge the assistance of Dr Pekka
Luhtanen in the mechanical calculation of the study.
Supported by a grant (No. 8318/78/78)from the Ministry
of Education (Finland).
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