Comments on Biomechanics and Muscle Coordination of Human Walking: Parts I
and II .
Zajac et al. [1,2], in their review of the biomechanics and muscle coordination of
human walking, encouraged the use of dynamical simulations to perform muscle-induced
segmental acceleration and power analyses to infer muscle coordination principles. They
state that a major function performed by a muscle arises from the instantaneous
segmental accelerations and redistribution of segmental energy throughout the body
caused by force generation, and this function can be fundamentally invariant to whether
the muscle is shortening, lengthening, or neither. Examples were provided, including
energy delivery to the crank by non-energy producing muscles in pedaling and forward
acceleration of the trunk (referred to as forward progression in Zajac et al. [2]) by
eccentric muscle activity in walking.
Although Zajac et al. [1,2] present a reasonable interpretation of muscle function
during locomotion using induced acceleration analyses of multi-linked, muscle-based
models, I feel that some of their interpretations should be placed more strongly in the
context of understanding established in simple, non-muscle-based models (e.g, inverted
pendulum model in walking) and muscle-based models with fewer degrees of freedom
(e.g., ankle-locked model in pedaling). The interpretations drawn from such simplified
models do not presume that muscular activity is not needed at the joints modeled as
constraints, but that muscle forces acting at these joints primarily provide postural
support. Moreover, seemingly novel concepts, such as a muscle s role in redistributing
segmental energy, can map fairly well into concepts revealed by these simplified models.
For example, in the analysis of pedaling in Zajac et al. [1], the interpretation that the
plantarflexors redistribute energy from the leg to the crank is not fundamentally different
from the simpler view that the plantarflexors lock the ankle, so that the proximal muscles
at the hip and knee accelerate the crank. In their interpretation, the instantaneous
cancellation of energy distributed to and from the leg, as energy is delivered to the crank,
does not appear to be physically relevant in differentiating between these two
mechanically-equivalent descriptions of energy delivery. Moreover, in tasks that are not
as well-understood as pedaling, the concept that a muscle simultaneously accelerates and
decelerates segments (i.e., redistribute energy) when it fundamentally supports the
posture of a joint can be misleading, since energy redistribution by the muscle may not
contribute importantly to net changes in individual segmental energy. In these instances,
relating the results from induced acceleration analysis of multi-linked models to
understanding established in simpler models can be particularly helpful.
For example, in the analysis of walking in Zajac et al. [2], the interpretation that
the proximal muscles that support the hip and knee (i.e., the vasti group and gluteus
maximus) contribute importantly to forward acceleration of the trunk by decelerating the
leg during the first half of stance (0-30% of gait cycle) does not seem appropriate. Based
on understanding derived from an inverted pendulum model, muscles that provide
postural support of the leg during the first half of stance are expected to lift and
decelerate the forward motion of the trunk. In agreement with this notion, the trunk is
observed experimentally to decelerate for much of this interval, and the ground reaction
force from the leg is directed backward, decelerating the center of mass. The question
that we should be asking is, If the leg doesn t function to accelerate the trunk forward
during the first half of stance, is it appropriate to emphasize the important contribution of
the proximal muscles to forward acceleration?
From about 15-30% of the gait cycle, the uni- and biarticular plantarflexors were
found to accelerate the trunk backward more strongly than the proximal muscles
accelerate it forward (see Fig. 3 in part II of the review) [2]. Therefore, during this
interval, the net effect of stance limb muscles (i.e., the proximal muscles and
plantarflexors) support and lift the trunk and decelerate its forward motion, which is
consistent with an inverted pendulum model of walking. However, Zajac et al. [2] chose
to emphasize the important individual contributions to forward acceleration provided by
the proximal muscles, as if to imply that accelerating the trunk forward during this
interval was necessary to maintain its forward motion.
From about 0-15% of the gait cycle, the presentation in Zajac et al. [2] that
proximal muscles at the hip and knee contribute importantly to forward acceleration is
even more perplexing. Other muscles do not appear to decelerate the trunk during this
interval (see Fig. 3 in part II of the review) [2], and the contributions of other, non-
muscular forces (i.e., passive torques, velocity effects, or gravitational forces), which
might explain the net effect of the leg, were not shown. Assuming that non-muscular
forces decelerate the trunk and cancel the forward acceleration contributed by muscles
during this interval, it would still seem inappropriate to emphasize the important
individual contributions of muscles to forward acceleration, when the net effect of the leg
decelerates the trunk, while it continues to progress forward through its momentum.
I believe that the energy redistribution from the leg to the trunk by the proximal
muscles during the first half of stance result indirectly from postural support, since they
cancel instantaneously with energy distributed away from the trunk by other forces. The
energy redistribution should not be interpreted to contribute importantly to forward
acceleration of the trunk to fulfill presumed task requirements of progression. These
issues have been communicated to Dr. Zajac and, hopefully, will be addressed in future
work.
Currently, the interpretation of muscle function during locomotion using induced
acceleration analysis is in its infancy, and its ultimate utility may depend on a better
understanding of the nature of the results and of the strengths and limitations of the
technique. As a first step, I feel that an appreciation of the mapping between
interpretations drawn from simple and complex models is needed before induced
acceleration analysis can be expected to advance our understanding of locomotion.
Indeed, the concept of synergistic muscle action in Zajac et al. [1], in which the net
accelerations induced by a group of co-excited muscles are interpreted to act as a
functional unit, can help clarify some of the important relationships between
interpretations drawn from different models of a task. I hope this commentary encourages
further debate between biomechanists and clinicians interested in the use of induced
acceleration analysis.
References
[1] Zajac FE, Neptune RR, Kautz SA. Biomechanics and muscle coordination of
human walking. Part I: Introduction to concepts, power transfer, dynamics and
simulations. Gait Posture 2002; 16: 215-232.
[2] Zajac FE, Neptune RR, Kautz SA. Biomechanics and muscle coordination of
human walking. Part II: Lessons from dynamical simulations and clinical
implications. Gait Posture 2003; 17: 1-17.
George Chen
Rehabilitation R&D Center, VA Palo Alto HCS,
Palo Alto, CA, USA and
Department of Mechanical Engineering,
Stanford University, Stanford, CA, USA
E-mail address: ****@*******.********.***
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