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Abstract
It has been stated repeatedly for the past 50 years that the steady-state force depression following shortening of an activated
muscle depends on the speed of shortening. However, these statements were based on results from experiments in which muscles
were shortened at different speeds but identical activation levels. Therefore, the force during shortening was changed in accordance
with the force–velocity relationship of muscles: that is, increasing speeds of shortening were associated with decreasing forces, and
vice versa. Consequently, it is not possible at present to distinguish whether force depression is caused by the changes in speed, as
frequently stated, or the associated changes in force, or both. The purpose of this study was to test if force depression depends on the
speed of shortening. We hypothesized that force depression was dependent on the force but not the speed of contraction. Our
prediction is that the amount of force depression after shortening contractions at different speeds could be similar if the force during
contraction was controlled at a similar level. Cat soleus muscles (n ¼ 7) were shortened by 9 or 12 mm at speeds of 3, 9, and 27 mm/s,
first with a constant activation during shortening (30 Hz), then with activation levels that were reduced (o30 Hz) for the slow speeds
(3 and 9 mm/s) to approximate the shortening forces of the fast speed contractions (27 mm/s). If done properly, force depression
could be precisely matched at the three different speeds, indicating that force depression was related to the force during the
shortening contraction but not to the speed. However, in order to match force depression, the forces during shortening had to be
systematically greater for the slow compared to the fast speeds of shortening, suggesting that force depression also depends on the
level of activation, as force depression at constant activation levels can only be matched if the force during shortening, evaluated by
the mechanical work, is identical. Therefore, we conclude that force depression depends on the force and activation level during
shortening, but does not depend on the speed of shortening as has been assumed for half a century. These results support, but do not
prove, the current hypothesis that force depression is caused by a stress-related cross-bridge inhibition in the actin–myosin overlap
zone that is newly formed during muscle shortening.
r 2004 Elsevier Ltd. All rights reserved.
0021-9290/$ - see front matter r 2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jbiomech.2004.09.028
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T.R. Leonard, W. Herzog / Journal of Biomechanics 38 (2005) 2190–2197 2191
2. Methods
Table 1
Mean force depression as a percentage of the isometric reference force
for trials at three speeds of shortening; 3, 9 and 27 mm/s
Activation during shortening was the same for all trials (30 Hz), and
the magnitude of shortening was the same for a given muscle (either 9
or 12 mm). Force depression decreases with increasing speeds of
shortening (po0:05) for these experimental conditions in agreement
with published findings.
3. Results
the same (i.e. within 0.2 N) were used for analysis. These
matched pairs were then used to calculate the work 3.2. Second test protocol
performed during the shortening phase by integrating
the force-shortening part of the test contractions In the seven muscles, 26 matched pair tests were
(Herzog et al., 2000). As a single measure of the force found for which the steady-state force depression
of shortening, the minimum force at the end of the was the same (i.e. within 0.2 N—Fig. 6) for two different
shortening phase was also used for the matched pair speeds of shortening. A total of 74 trials where
analysis. the stimulation frequency was manipulated were per-
Statistical analysis was performed using the Wilcoxon formed to obtain the 26 successfully matched pairs.
signed rank test (Stata software, Stata Corp., College Twenty-six matched pair tests were deemed sufficient
Station, TX, USA) to test for differences between the to proceed with further data analysis due to the
force depressions of matched trials to determine whether high quality of the data traces obtained. Statistical
the trials were truly matched. The sign test was used to analysis revealed that force depression in the slow and
test for differences in force or work performed for fast speed trials was the same (p ¼ 0:708), that means
the matched test trials, that is, all trials for which there was no consistent bias towards any of the
the speed of shortening was different but force depres- experimental speeds.
sion was the same. The null hypothesis was that the However, the minimum force during shortening and
speed of shortening does not influence force depression. the work performed during shortening was always
In other words, it was expected that for the matched greater in the slow compared to the fast speed for each
pair trials, the minimum force during shortening and matched pair (po0:05; Figs. 6 and 7). Therefore, the
the work performed during shortening were the same. null hypothesis that force and work during shortening
All statistical analyses were performed at a level of would be the same for tests with the same steady-state
significance of a ¼ 0:05: force depression was rejected.
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T.R. Leonard, W. Herzog / Journal of Biomechanics 38 (2005) 2190–2197 2195
Fig. 7. Box and whisker plot showing median, 25th percentile and 75th percentile values for parameter differences for all force depression matched
pairs (n ¼ 26). Differences were calculated as follows: test parameter (force depression (a) or minimum force (b) or work performed (c)) for the slow
speed of shortening minus the test parameter for the faster speed of shortening. Note that the median difference in force depression for the matched
pairs (a) is close to zero and the whiskers extend into both positive and negative zones. Differences for both work production (b) and minimum force
(c) were positive for all 26 independent observations.
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2196 T.R. Leonard, W. Herzog / Journal of Biomechanics 38 (2005) 2190–2197
to evaluate this expectation, we quantified the minimum active muscle can be the same for different speeds of
force at the end of the shortening phase as a single force shortening if force is adjusted appropriately. These
indicator, and also determined the work performed by results strongly suggest that force depression does not
soleus during shortening as a global indicator of the depend on the speed of shortening as stated throughout
force profile. We found that the minimum force during the literature of the past 50 years, but that force
shortening, as well as the work performed was always depression depends directly on the force during short-
greater in the matched pair trials for the slow compared ening, and that the association with speed is merely an
to the fast speed (i.e. always greater for 3 mm/s artifact of the force–velocity relationship which causes
compared to 9 or 27 mm/s, and always greater for 9 changes in force during the shortening phase when
compared to 27 mm/s; Fig. 7). Therefore, aside from the experiments are performed at different speeds of short-
force during shortening, it appears that another factor ening (Hill, 1938). Therefore, in the search for mechan-
plays a role in force depression, albeit a small one. The isms, force should be considered whereas speed does not
present experiment was not designed to determine the appear to have an effect on force depression.
origin of this other effect. However, we would like to
tentatively suggest that it is associated with the amount
of activation and not the speed of shortening. We
Acknowledgements
eliminate the speed of shortening as a candidate because
there is a very tight relationship between the work
Funding for this project was provided by the Natural
performed by a muscle and the amount of force Sciences and Engineering Research Council of Canada
depression for tests at different speeds but constant (NSERC).
activation (Herzog et al., 2000). This relationship does
not hold here at all as the same force depression is
achieved for vastly different amounts of mechanical References
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