Tg. Sandercock et Cj. Heckman, FORCE FROM CAT SOLEUS MUSCLE DURING IMPOSED LOCOMOTOR-LIKE MOVEMENTS - EXPERIMENTAL-DATA VERSUS HILL-TYPE MODEL PREDICTIONS, Journal of neurophysiology, 77(3), 1997, pp. 1538-1552
Muscle is usually studied under nonphysiological conditions, such as t
etanic stimulation or isovelocity movements, conditions selected to is
olate specific properties or mechanisms in muscle. The purpose of this
study was to measure the function of cat soleus muscle during physiol
ogical conditions, specifically a simulation of a single speed of slow
walking, to determine whether the resulting force could be accurately
represented by a Hill-type model. Because Hill-type models do not inc
lude history-dependent muscle properties or interactions among propert
ies,the magnitudes of errors in predicted forces were expected to reve
al whether these phenomena play important roles in the physiological c
onditions of this locomotor pattern. The natural locomotor length patt
ern during slow walking, and the action potential train for a low-thre
shold motor unit during slow walking, were obtained from the literatur
e. The whole soleus muscle was synchronously stimulated with the locom
otor pulse train while a muscle puller imposed the locomotor movement.
The experimental results were similar to force measured via buckle tr
ansducer in freely walking animals. A Hill-type model was used to simu
late the locomotor force. In a separate set of experiments. the parame
ters needed for a Hill-type model (force-velocity, length-tension, and
stiffness of the series elastic element) were measured from the same
muscle. Activation was determined by inverse computation of an isometr
ic contraction with the use of the same locomotor stimulus pattern. Du
ring the stimulus train, the Hill-type model fit the locomotor data fa
irly well, with errors <10% of maximal tetanic tension. A substantial
error occurred during the relaxation phase. The model overestimated fo
rce by similar to 30% of maximal tetanic tension. A nonlinear series e
lastic element had little influence on the force predicted by a Hill m
odel, yet dramatically altered the predicted muscle fiber lengths. Fur
ther experiments and modeling were performed to determine the source o
f errors in the Hill-type model. Isovelocity ramps were constructed to
pass through a. selected point in the locomotor movement with the sam
e velocity and muscle length. The muscle was stimulated with the same
locomotor pulse train. The largest errors again occurred during the re
laxation phase following completion of the stimulus. Stretch during st
imulation caused the Hill model to underestimate the relaxation force.
Shortening movements during stimulation caused the Hill model to over
estimate the relaxation force. These errors may be attributed to the e
ffects of movement on crossbridge persistence, and/or the changing aff
inity of troponin for calcium between bound and unbound crossbridges,
neither of which is well represented in a Hill model. Other sources of
error are discussed. The model presented represents the limit of accu
racy of a basic Hill-type model applied to cat soleus. The model had e
very advantage: the parameters were measured from the same muscle for
which the locomotion was simulated and errors that could arise in the
estimation of activation dynamics were avoided by inverse calculation.
The accuracy might be improved by compensating for the apparent effec
ts of velocity and length on activation. Further studies are required
to determine to what degree these conclusions can be generalized to ot
her movements and muscles.