Acta physiol. scand. 1976. 96.

83-93
From the Laboratory for the Theory of Gymnastics, August Krogh Institute,
University of Copenhagen, Denmark

Mechano-Elastic Properties of Human Muscles at

Different Temperatures
BY
ERL~NG FLEMM~NG
ASMUSSEN, BONDE-PETERSEN
and KURTJ0RGENSEN

Received 24 June 1975

Abstract
ASMUSSEN,E., F. BONDE-PETERSEN and K. JBRGENSEN. Mechano-elasticproperties of human
muscles at different temperatures. Acta physiol. scand. 1976. 96. 83-93.
The effect of changes in the muscle temperature on their ability to store elastic energy was studied by hav-
ing 5 trained subjects perform maximal vertical jumps on a force platform, with and without counter move-
ment, at muscle temperatures between about 32°C and 37°C. The results showed that the heights of vertical
jumps were considerably reduced at lowered temperature, but the gain in heiglir after a counter movement
in the form of a jump down from a height of 0.4 m over the force platform, was significantly higher in the
cold condition. T o test whether this was due to an increased stiffness of the muscles, experiments with im-
posed sinusoidal length variations at 14 Hz were performed. Aforce x Alength-L (i.e. stiffness) increased
with isometric tension independent of muscle temperature. Experiments in which the rate of tension de-
velopment and relaxation in voluntary maximal isometric contractions were measured at different muscle
temperatures showed that maximal isometric tension changed by less than I X per degree but the rate of
tension development and relaxation by 3-57, and 5 % per degree, respectively, in the temperature range
studied (30” to 40’). These data may be explained by the hypothesis that the series elastic components of
the active muscle are located in the cross-bridges between myosin and actin filaments. The storage of elastic
energy would be enhanced if the rate of breaking of these bridges were decreased at lower temperatures.
Key words: Elastic energy: muscle stiffness; muscle temperature; rate of tension development and relaxa-
tion; vertical jumps

Recently we demonstrated in man that mechanical energy imparted to a muscle while

it is actively resisting lengthening, can be stored and reused as elastic energy during a sub-
sequent shortening contraction (Asmussen and Bonde-Petersen 1974). This happens in a
vertical jump when the jump is preceded by a countermovement. We assumed that the ac-
tive muscles behaved as springs, and found that an average of up to 23% of the imparted
mechanical energy could be stored as elastic energy and re-used in the subsequent jump.
Marey and Derneny (1885) and Cavagna et al. (1971) came to similar conclusions. The na-
ture of the elastic component that allows active muscles to store energy is not fully known,
nor is its location. The expressions “parallel-elastic” and “series-elastic” were proposed by
Levin and Wyman (1927) to designate the elastic components in parallel with or in series
84 ERLING ASMUSSEN, FLEMMING BONDE-PETERSEN AND KURT JBRGENSEN

with the contractile elements in the muscle. The former is usually identified as tendons, intra-
muscular connective tissue, sarcolemma etc., whereas the latter, in recent literature, has
been assumed to be in the cross-bridges that form between myosin and actin filaments
during contraction (A. F. Huxley and Simmons 1971, Riiegg 1971, Rack and Westbury
1974, A. F. Huxley 1974). With these possibilities it seemed relevant to study the effect of
temperature on the elastic properties, including storage of elastic energy, in human muscles
in situ. As a working hypothesis we assumed that lowering of the muscle temperature would
increase the stiffness of the visco-elastic components while at the same time decreasing the
rate of the chemical processes that underlie the formation and breaking of the cross-bridges.

Methods
Muscle reniperarurcs. Variations of muscle temperature were obtained by immersing the subject in cold
water, or by active exercise. The muscle temperature was measured by means of a thermo needle (Ellab)
inserted into the appropriate muscle to a fixed depth of 25-30 mm. There was a rather steep temperature
gradient from the surface inward, especially after cooling. The temperatures measured, therefore, do not
represent mean muscle temperatures but are arbitrary values for comparisons between cold and warm
muscles.
Slorage of elastic energy in the muscles of the legs was estimated from the increase in height of a vertical
jump performed in direct continuation of a jump down from a height of 0.40 m, over the height when i t
was performed from a semi-squatting position (cf. Asmussen and Bonde-Petersen 1974). The jumps were
performed on a force platform (Bonde-Petersen 1975) in connection with an inkwriting recorder at a paper
speed of 125 mm x s-'. The registered time, tnlght. between take-off to the jump and touch-down after the
jump was used in calculation of h, the height of the jump, according to the formula

h = vws2 (2g)-'
>
. (1)

Here, g is the acceleration of gravity (9.81 m 4 s-~) and vpo8 is the velocity at take-off (and touch-down).
This velocity is determined from the time spent in moving upwards (or downwards), i tilight, as vpoB= 4
tfllghL g. The height of the jump consequently is h = ( i tilight ?: g)2x (2g)-'= 1.226 (tfIlghJd. When the
subject jumps down from an elevation onto the force platform before the actual upwards jump, a certain
time, ttotal, is spent in contact with the platform. Part of this time, tneg, is spent in absorbing the energy
from the fall down and the rest (ttotal-tneg) in liberating energy for the take-off. The braking movement
will take place with a downward velocity that changes from vneg (at first contact with platform) to zero,
and the subsequent upward movement, leading to the jump, will have velocity zero at the beginning of the
movement and end with velocity=v,,,, as calculated above. The distance covered in the decelerating and
accelerating movements, respectively, must be the same; hence from the mean velocities, tnep y 4 vncg-
(ttotal-tnes) x 4 vpoB.Solving for tnCgone gets

tneg = ttotsl vpos (Vneg+ vpos)-* (2)

In this way the time during which energy is absorbed can be calculated. The rest of the contact time with
the platform is used in liberating positive kinetic energy.
Muscle sfqfness. An arbitrary expression for the stiffness (Aforce/Alength) of the calf muscles was ob-
tained by imposing sinusoidal length variations of constant amplitude and frequencey on the isometrically
contracted muscles, registering the resulting rhythmic variations in force around the mean muscle tensions.
For this purpose the subject lay o n his back, his flexed knee pressed against a solid cross-b:am and the ball
of his foot lightly tied to a wooden sandal, movably fixed to a ring-shaped strain gauge dynamometer (see
Fig. 1). The subject was asked t o press the ball of his foot against the dynamometer by a plantar flexion
with a pre-determined force and to keep this constant by watching the hand of the strain gauge meter. The
hand was so much damped that it could not follow the rapid sinusoidal oscillations induced by an ec-
centric driven by a strong motor (see Fig. 2). We found that 14 H z gave good and reproducible force varia-
tions with movements of 40 mm peak-to-peak. The length variations at the Achilles tendon must have been
 MECHANO-ELASTIC PROPERTIES OF MUSCLE 85

Fig. I . Schematic drawing of set-up for vibrational

studies of m.triceps surae. a. wall, floor. 6. support-
ing frame. c. ring-shaped strain gauge dynamometer.
d. wooden sandal, movably attached to r. e. solid
cross beam. f. iron lever, movably attached to 6. g.
motor and eccentric.

about one-third of this, 12-14 mm. Each period, at a maintained constant mean force, lasted 2 to 3 s. Emg
mean peak voltage was registered simultaneously by means of skin electrodes over the soleus muscle, both
before and during the vibration. Muscle temperatures were measured in the lateral head of the gastro-
cnemius muscle at a depth of 25 mm.
Rare of tension delfoelopment and relaxation.' The speed of these two parameters was measured during
maximal voluntary isometric contraction of the elbow flexors, the plantar flexors, and the knee extensors-
henceforth called, respectively, biceps br., triceps surae, and quadriceps fem. The registration of the iso-
metric tensions was performed by means of a strain gauge dynamometer with the elbow, ankle, or kneein
a standard position. Emg and mean peak voltage were registered simultaneously from double surface
electrodes by a Disa electromyograph and an inkwriting recorder (Mingograph) at a paper speed of 250
mm s-I.
The maximum mechanical tensions in the cold or warm conditions were expressed as percentages of
the maximum isometric tensions at normal temperature o n the same day. Because of the uncertainty of
defining the exact beginning and end of the tension development, the rate was expressed as tension increase
from 5 t o 90% of maximum over time (Fig. 3). Correspondingly, the speed of relaxation was calculated
from the time between 905: and 5:: of maximal tension during relaxation. The rates of tension develop-
ment and relaxation were expressed as percentages of the controls. In this way all three muscle groups,
from three different subjects, could be combined in the same graph with the measured temperature as
abscissa (Fig. 6-8).

Fig. 2. Examples of vibrational

ustr.
900

600
1 ustr.
900

600
force variations at increasing mean
tensions in m.triceps surae. Length
3 00 300
variations at Achilles tendon:

-
about 12 mm, vibrational fre-
quency 14 Hz, arbitrary force 0 - 0
units in jt-strain.

1 sec

The experiments in this section were performed by Mr. Finn Andersen as part of his thesis for the cand.
scient. degree.
86 ERLING ASMUSSEN, FLEMMING BONDE-PETERSEN AND KURT J0RGENSEN

(0.05 - 0.9)P. Fig. 3. Lower curve: tension de-

velopment in maximal voluntary
isometric contraction of m.triceps
surae. Upper curves, iemg (mean
,~ peak voltage) and emg. Rate of
tension development is taken t o
be P0(0.9- 0.05) x t-I where Po is
maximal tension and t time in
I_
seconds.
50 H r

Subjects were 8 young men, students of physical education, of good health, and well fit for muscular
exercise. They did not all serve in all the experiments t o be presented, but each experimental situation was
repeated several to many times o n a given subject.

Results
Vertical jumps, from a squatting position or in continuation of a jump down from a height
of 0.40 m, were performed at different temperatures of the leg muscles. The heights of the
jumps, calculated as described under Methods, were plotted against the muscle tempera-
tures, measured in the lateral vastus muscle at a depth of 30 rnm, as shown in Fig. 4 for one
of the 5 subjects. It is evident that the height of a vertical jump in both cases is smaller in
the cold condition. But it also came out that the gain in height due to the effect of jumping
down 0.40 m, was larger both absolutely and relatively in the cold condition. Assuming

Height of jump
0.5

0.L '

0.3 . Fig. 4. Ordinate: The height of a

vertical jump performed from the
semisquatting position ( 0 ) .or in
continuation of a jump from
height 0.40 m ( A ) . Abscissa:
Temperature in m.vastus lat. at a
L-
29
I

30
I

31
.
32
1

33
I

31
,
35
,
36
,
37
,
38
,
39
,
LO 'C
, depth of 30 mm. Subject M .
 MECHANO-ELASTIC PROPERTIES OF MUSCLE 87

1. y = a + bx.
TABLE

Subject Squatting jump After jump down 0.4 m

a b r a b r

F. -0.929 0.037 0.938 -0.668 0.030 0.920

A. -0.854 0.034 0.945 -0.695 0.031 0.945
S. -0.557 0.024 0.978 -0.444 0.021 0.973
M. -0.528 0.024 0.907 -0.258 0.018 0.837
B. -0.531 0.025 0.905 -0.326 0.021 0.894

Constants in formula y = a + bx, in which y = height of jump in m, and x is muscle temperature in C". r =
correlation factor.

the values to lie on straight lines (see Fig. 4), the regression equation for the two kinds of
jump was calculated as y = a + bx, where y is the height of the jump, x is the muscle tem-
perature, and a and b are constants. These constants, and the r-values for the correlation,
are presented in Table I.
If all data from a warm condition (about 37°C) are treated as one series of experiments,
and all data from the corresponding cold condition (about 32°C) as another series, average
height of the jumps with and without the preceding jump down, can be calculated for each
subject. Further, the average gain in height, Ah, after jumping down, over the height ob-
tained in the squatting jump, can be calculated for both series. These data are presented in
Table 11. The table shows that in all five subjects the gain in jumping height after a jump
down, Ah, was larger with cold muscles than with warm muscles. The average Ah-difference
was significant at the 0.01 level, and the individual differences were significant at the 0.1
level in all 5 subjects and at the 0.05 level or better in 4 of the 5 subjects (Table 11).
The absorption and storage of energy from the down-jump takes place during the first
part of the time spent in contact with the force platform. Its duration, tnEg, can be calcu-
lated-as shown under Methods-from formula (2). Of the parameters in the formula vpos
is calculated from formula (1) in which h is determined from the appropriate regression
equation (Table I) for muscle temperatures 37°C and 32"C, respectively. vnEgis a constant,

TABLE
11.

Gain in height after jumping down

Subject Warm conditions Cold conditions Warm-cold

difference
n Tkus ih+S.E. n T&S Ah&S.E.
C" m C" m

F. 11 37.1 0.012f0.013 II 33.1 0.039f0.006 0.05 P--0.1

A. I2 36.5 0.037f0.005 12 32.2 0.058F0.005 P :c 0.01
S. 16 37.5 0.005+0.003 7 33.2 0.020f0.003 P~O.001
M. 10 37.7 0.018f0.006 9 32.9 0.055 f 0.005 P<O.OOI
B. 7 39.6 0.017k0.008 7 32.3 0.052k0.012 P .' 0.05
Mean 0 .0 1 7 0 ~0 .0 0 7 5 46 0.0462k 0.0066 P < 0.01
88 ERLING ASMUSSEN, FLEMMING BONDE-PETERSEN AND KURT J0RGENSEN

TABLE
111.

Subject Body After jump down, 0.40 m

mass
kg Warm conditions (37°C) Cold conditions (32°C)

h vpos [tot tneg tpos h vpos ttot tneg tpos

m m/s s S S rn m/s s S S

F. 69 0.442 2.94 0.249 0.128 0.121 0.292 2.39 0.264 0.126 0.138
A. 94 0.452 2.98 0.289 0.149 0.140 0.297 2.41 0.326 0.151 0.175
S. 64 0.333 2.56 0.256 0.122 0.134 0.228 2.12 0.272 0.117 0.155
M. 75.5 0.408 2.83 0.196 0.099 0.097 0.318 2.50 0.217 0.102 0.115
B. 72 0.451 2.97 0.201 0.103 0.098 0.346 2.61 0.223 0.107 0.116
Mean 74.9 2.86 0.120 0.118 2.41 0.121 0.140

depending only on the height, 0.40 m, of the elevated platform from which the down-jump
takes place and hence is vnes =0.40 x 2g =2.80 m x s-l. The total time in contact with the
force platform, ttotal, is measured and averaged for each subject at temperatures about
37°C and 32°C-neglecting the small variations around these temperatures in the actual
mean values. Individual mean values of these parameters and the resulting values of tneg
and tpos - (ttotal- tneg)are presented in Table 111.
It appears that whereas tnep is uninfluenced by lowering of the temperature, tpoa is longer
in the cold condition. From the constancy of tnegand vneg in both conditions, it follows that
the distance covered during the negative phase is constant-presumably because it is steered
voluntarily by the subject. In spite of the longer time available for positive work in the cold
condition, the final velocity, vDos is smaller. That implies that the mean resulting force, P,
developed during this time, P = m x vpos x tpoB-l is smaller in the cold condition than in
the warm condition (1289N and 181SN,respectively).
The actual force developed by the muscles and acting on the platform must be larger by
the amount of gravitational force on the subjects-on an average by 74.9 ' 9.81 735N.
The ratio between these maximal forces thus becomes (1 289 I 735) ( I 815 +735)-' 0.76,
A

range 0.73 to 0.81.

In the squatting jump, where no countermovement is performed tneg consequently is 0,
and hence tDosis equal to ttota,. The tposwas not significantly different in the two conditions,
but the resulting force, P, and the total force, (P +weight), was also smaller here in the cold
condition ( I 223N) than in the warm condition (1 399N). The ratio between the total forces
was 0.87 on an average, the range being 0.86 to 0.90 for the 5 subjects (cf. Table IV).
Muscle stiffness. Fig. 2 shows examples of the force variations recorded during sinusoidal
length variations at 14 Hz applied to the forefoot during isometric plantar flexions of vari-
ous tensions. The peak-to-peak changes in p-strains are used as arbitrary expressions for
the stiffness of the calf muscles. When averaged from 4 to 5 expts on the same subject and
plotted against the mean force in the muscle group, stiffness was found to increase with
voluntary isometric force as shown in Fig. 5 for one subject. The shape of the curves could
vary somewhat from subject to subject, being concave (as in Fig. 5 ) or closer to a straight
line. When curves from experiments performed after cooling of the leg muscles to 3&32 'C
 MECHANO-ELASTIC PROPERTIES OF MUSCLE 89

I
L:
amplitude f SE
#strain
500

400 r 1
Fig. 5. vibrations
during
m.tricepsAverage
surae atforce
different
at 14 Hz
variations
mean
of 300

tensions (abscissa). :.: - x at

3 7 T , 0-0 at 31°C muscle tem- ZOO
perature. Each point is mean of
5 determinations. Vertical bars
den0tekS.E. Maximum volun- loo
tary tension I030 p strain in
both conditions (not shown). , mean tension
0
100 200 300 400 500 600 700 800 #strain

were superimposed on the curves at normal muscle temperatures they very nearly coincided
(as in Fig. 5). Only in one out of five subjects was the curve from the cold experiment sys-
tematically, but statistically insignificantly, displaced 10-1 5 7,upwards. The stiffness at any
force thus seems to be uninfluenced by the temperature. The maximum isometric force was
in all subjects uninfluenced by the temperature shift.
Mean peak voltage, from emgs picked up by skin electrodes over m.soleus, increased
almost linearly with mechanical force. But here it was found that for a given force the mean
peak voltage was always higher at the low temperature (3e32"C) than at normal tempera-
ture (37-38°C).
The maximum isometric tension, Po,measured in biceps br., triceps surae, and quadri-
ceps fern., in relation to muscle temperature, is shown in Fig. 6 for 3 subjects. In absolute
values the mean maximum isometric tension, measured at the wrist (biceps br.), at the ball
of the foot (triceps surae), and just above the malleoli (quadriceps fern.) were for the 3 sub-
jects 33 kp (324N)+7% (SD), 92 kp (902N)k10% (SD), and 50 kp (491N)+7% (SD).
Assuming all values to lie around a straight line, the regression equation becomes y =
68.4 +0.81x with an r value of 0.32 (P-- 0.001). It shows that lowering the temperature from
40" to 3OoC, lowers Poby only 8%.
Fig. 7 shows the effect of lowering the muscle temperature on the percentage rate of
tension development from 5 to 90% of maximum (Fig. 3). The actual rates at normal tem-

4 Y. of conlrol
max. i s o m . force

Fig. 6. Maximum voluntary iso-

metric tension (Po) at different
muscle temperatures in per cent
140
120 i x o .. .
of daily control value. Data from
three muscle groups (different
signatures): mtriceos brachii. m.
lo
. ' *i
.8.

triceps surae, and m.quadriceps

fem., on each of 3 subjects.
27 28 29 30 31.. 32 33 34 35 36 37 38 39 40 II Co
90 ERLING ASMUSSEN, FLEMMINO BONDE-PETERSEN AND KURT JQRGENSEN

r a t e of lension devclopmcnl

140

ment, (0.05 Po to 0.9 Po) in per

cent of controls in relation to
muscle temperature. 3 muscle
groups, in 3 subjects. Explanation

perature were 4.75 Pox s-l for biceps br., 2.31 Po x s-l for triceps surae, and 5.00 Po s-I
for quadriceps fem. Again assuming the points to scatter around a straight line, the regres-
sion becomes y = - 33.3 + 3.45x, with r =0.59 (P <0.001). Thus a temperature drop of 10°C
will cause a drop in speed of tension development of about 35 %.
Fig. 8 shows the effect of temperature change on the percentage rate of relaxation, from
90% to 5 % of Po. The actual normal average values for the three muscles were 3.30 Po \I

s-' (biceps br.), 1.15 Po x s-' (triceps surae), and 2.35 Po s-l (quadriceps fern.). The straight
line regression will be y = -89.4 + 5.03x, r =0.79 (PiO.001). A drop of temperature of 10°C
will cause a decrease in the speed of relaxation of about 50%.
These large temperature dependent changes in rates of tension development and relaxa-
tion were not caused by changes in the speed of neuro-muscular activation as revealed by
the emg; as seen in Fig. 3, the full mean peak voltage has been reached before 5 % Po is
reached.

Discussion
The results from the squatting high jumps in Table I and Fig. 4 show that there is a strong
positive correlation between the muscle temperature and the height of the jump. The slopes
of the individual regression lines (b in Table I) are different in the 5 subjects, but it must be
remembered that the muscle temperature as measured is not an average o r weighted muscle
temperature but an arbitrary sample of the temperature inside one of the active muscles.
The height of the vertical jump after a jump down (Table I, Fig. 4) is also strongly cor-
related to the muscle temperature, and the slopes of the regression lines (b in Table I) are
again individually different. What is noteworthy is that these slopes are always lower after
a jump down than in the squatting jump. Fig. 4 demonstrates this for one subject, and it is
obvious that the difference in the heights of jumps performed with or without the initial
jump down is larger at low temperatures than at normal or higher temperatures, where the
difference may dwindle towards zero. The gain in height of a vertical jump which can be
obtained after a preliminary countermovement or jump down has been ascribed to the stor-
age of elastic energy in the muscles in the period where they are being stretched (Marey and
 MECHANO-ELASTIC PROPERTIES OF MUSCLE 91

180

I60
I
140.
7. of control
r a t e of relaxallon

Fig. 8. Rate of relaxation (0.9 Po .Ct; "'* *

t o 0.0sPo)in relation to muscle
temperature. Explanation as in 20:
Fig. 6. ,

Demeny 1885, Cavagna et al., 1971. Asmussen and Bonde-Petersen 1974). The present ex-
periments accordingly demonstrate that more elastic energy can be stored in cold muscles
than in warm muscles. This difference in elastic energy storage, as visualized by the greater
gain of height, Ah, at low temperatures, is significant at the 0.01 level (Table 11).
The preliminary jump down was in all cases performed from a height of 0.40 m. The
energy from which the elastic energy store could be drawn thus was the same in both cold
and warm conditions. Further, the time used in absorbing this energy, tnep,appeared t o be
uninfluenced by temperature, although individually different (Table 111). It follows, as
calculated in Results and shown in Table 111, that at the turning point from a downward
directed to an upward directed movement of the body's center of gravity, the same amount
of energy has been absorbed, during the same time, and over the same distance, in cold
and warm conditions. Nevertheless more of this energy apparently can be stored and re-
used in the actual jump when this is performed with cold muscles than when performed with
warm muscles. With warm muscles a gain in height of from 0.005 to 0.037 m (average
0.0178 m) was achieved, but with cold muscles the gain was from 0.020 to 0.058 m (average
0.0448 m) (Table 11). As the jump down in all cases was 0.40 m, the re-used energy becomes
4.45% of the absorbed energy (range 1.25-9.25%)in the warm condition, but 11.2%(range
5.0-14.5%) in the cold condition.
An increased ability for storing elastic energy would obtain if the elastic stiffness of the
muscles (Aforce/Alength) had increased. The present studies over the force-variations
caused by sinusoidal length variations in warm and cold conditions showed, however, that
referred to the same mean muscle tension, the stiffness of the muscles was not measurably
influenced by temperature changes in the range 30°C to 37°C (Fig. 5 ) . This was unexpected
because generally tissues like collagenous tendons (Apter 1972), joints (Rice 1967), and
muscle fibers (Buchthal and Kaiser 1951) become stiffer at lower temperatures. But if muscle
elasticity essentially is located in the active cross-bridges of the muscle filaments (Huxley
and Simmons 1971, Rack and Westbury 1974, A. F. Huxley 1974), it seems natural that at
the same mean tension, when the same number of cross-bridges must be engaged, irrespec-
tive of temperature, the elastic stiffness must be much the same. (Whether the forced sinus-
oidal movements elicit stretch reflexes or not (cf. Joyce et al. (1974)) is unimportant here,
92 ERLING ASMUSSEN, FLEMMlNG BONDE-PETERSEN AND KURT JBRGENSEN

Subject Body mass Squatting jump

k6
Warm conditions (ca. 37°C) Cold conditions (ca. 32'C)
b
ttot ha Vp o s b ttot ha VPOS
s m mls S m mls

F. 69 0.28 I 0.440 2.94 0.268 0.255 2.10

A. 94 0.392 0.404 2.82 0.43 I 0.234 2.14
S. 64 0.378 0.331 2.55 0.372 0.21 I 2.03
M. 75.5 0.243 0.360 2.66 0.268 0.240 2.17
B. 72 0.332 0.394 2.78 0.349 0.269 2.30
Mean 74.9 0.315 0.386 2.75 0.338 0.242 2.15

a From regression in Table I.

From vpos= ]I=.

where the conscious subject continuously adjusts the mean force to the pre-determined
level.)
A possible explanation for the increased ability to store and re-use elastic energy in the
cold condition may be found in the fact that the formation and disengagement of cross-
bridges probably takes a longer time in the cold condition than at higher temperature.
This is supported by the longer time of both tension development and relaxation demon-
strated in Fig. 7 and 8. The slower tension development explains why the subjects at low
temperatures in the squatting jump were able to attain a height of only about 60c>:,of that
at the higher temperature. The distance through which the muscles could shorten at take-
off was-as mentioned under Results-identical in the two conditions. A slower develop-
ment of maximum force consequently leads to less work produced during the positive phase
of the jump, even though the time, ttot, is prolonged by about 7 % (Table IV), because the
muscles in this time apparently could reach an average tension of only 87 % of that reached
in the warm condition.
After the jump down from height 0.40 m the muscles are activated to d o negative work.
The average tension developed for this purpose was-as follows from the identical tnep
(Table Ill) and absorbed kinetic energy-identical in cold and warm conditions. As relaxa-
tion takes a longer time in the cold condition (Fig. 8), it will mean that the increased stiff-
ness of the active muscles disappear later and thus will be able to carry more elastic energy
over into the positive phase of the jump. This explanation is in good agreement with the ex-
periments in Fig. 7 and 8, and also with the finding that isometric endurance is prolonged at
lower temperatures (Clark, Hellon and Lind (1958), Edwards et a/. (1970)). The latter authors
also found that the ATP-fluxes, as determined from muscle biopsies of human muscles,
were considerably lowered at low muscle temperatures.
Our conclusion is that both the generally lower jumping height and the increased ability
to carry over energy from a jump down to a subsequent vertical jump can be explained by
assuming that the formation and breaking of active cross-bridges in contracting human
muscles is considerably delayed at lower temperatures, even though the final maximal ten-
sion is only slightly decreased.
 MECHANO-ELASTIC PROPERTIES OF MUSCLE 93

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