Abstract— In this article the principles of human locomotion
are revisited and reviewed. This has been done in the
framework of two European projects, where the elicitation of
these mechanisms inform, on the one hand, the design of
artificial bipedal walkers (H2R), and on the other hand the
design of lower limb exoskeletons (BETTER) for rehabilitation
of gait in post-stroke patients. Passive dynamics emerging from
the morphology of the human musculoskeletal system, reflexes
as stabilization mechanisms, modular control of movement as
well as supra-spinal control of gait are reviewed to get insight
on how these mechanisms can be used to explain human
locomotion.
I. INTRODUCTION
The human musculoskeletal and neural-motor system is
highly optimized for efficient locomotion. Efficiency,
stability and voluntary modulation of human gait are a result
of a combination of features spanning the human
musculoskeletal, sensorimotor and neural systems. Salient
aspects of these systems include (1) the functional
morphology, (2) the synergistic coordination of motor
activity, (3) the phase dependent modulation of muscle
activity and (4) cognitive skills.
On the one hand, the functional morphology is highly
optimized for efficient biped locomotion [1] as it allows
exploiting the inherent dynamics to reduce energy
consumption and control effort, and result in natural looking
motions. Also a contribution of this functional morphology is
the capability of self-stabilization, since the elastic properties
of muscles and tendons increase stability without active
control.
On the other hand, the synergistic feed-forward motor
patterns, be it activated at kinetic or kinematic events or due
to learned timing, create coordinated synergies of movement.
Whilst feedback control occurs at various levels of
complexity regarding the extension of perception and
deployment of muscle action, phase-dependent modulation is
a function of the current task or phase of motion, and as a
result reflex action can be modulated, reinforced, or
suppressed. Eventually, cognition plays a crucial role in
learning and predicting the sensory consequences of actions,
helping to deal with feedback time delays and allowing for
planning the appropriate compensative actions (active and
passive).
*Research has been supported by The Spanish CONSOLIDER Program
through Grant CSD2009-00067 and by the European Commission FP7
through grants 247935 (BETTER) and 600698 (H2R).
J.L. Pons (corresponding author, phone: +34918711900; fax:
+34918717050; e-mail: jose.pons@csic.es), J.C. Moreno and D. Torricelli
are with the Bioengineering Group, Spanish National Research Council,
CSIC, Arganda del Rey, Madrid, Spain.
J.S. Taylor is with the Hospital de Parapléjicos de Toledo, Toledo,
Spain.
This paper will address and review the various motor
principles leading to efficient locomotion in humans. In so
doing, we will analyze the functional morphology in section
II, the contribution of reflex mechanisms to stabilize
locomotion in section III, the orchestration and synergistic
coordination of motor patterns in section II and eventually
the supra-spinal control of gait in humans in section IV.
II. FUNCTIONAL MORPHOLOGY OF THE HUMAN
MUSCULOSKELETAL SYSTEM
Research on the biomechanics of human locomotion
provides valuable insights into basic principles for motor
control. Human legged locomotion is so efficient partially
because it does not power movements with independent
motor actions at each joint. Muscles often span multiple
joints, which results in energy-saving and power transfers
when a movement simultaneously requires negative power at
one joint and positive power at another joint [2]. This allows
making effective use of passive elastic properties to generate
part of the required force or power without metabolic cost,
especially when muscle-tendon units span multiple joints [3].
Furthermore, multiple muscles spanning a joint allow
efficient modulation of joint stiffness during dynamic
movements, making fast adaptation to uneven surfaces and
terrains possible.
Passive dynamic walking introduced, as a model, in
under-actuated bipedal walkers and robots shows emerging,
natural-looking walking gait with remarkable similarities to
human walking. In this regards, passive dynamic walking,
first introduced by McGeer [4], exploits the mechanical
potential energy gained while walking down a slope, showing
a stable gait without any or limited control or actuation. Limit
Cycle (LC) walking machines represent a step forward in this
direction. They combine the exploitation of passive dynamics
with minimal feed-forward actuation in order to replenish
energy losses and to increase stability. Examples of LC
walking prototypes have been developed by TU Delft [5],
Cornell University and MIT [6]. The Cornell and Delft
bipeds demonstrate that basic walking can be accomplished
with extremely simple control and very low energy
consumption. However, due to absence of feedback control,
passive dynamics bipeds cannot react to disturbances or
external forces, even though human-like gait is achieved,
walking is unstable as all other stabilization mechanisms
found in humans are still lacking.
III. ROLE OF REFLEX FUNCTION FOR STABILIZATION OF
LOCOMOTION
Bipedal plantigrade walking in humans is unique
compared to digitigrade locomotion in animals, in that the
stance phase is made by ground contact first by the heel, then
the foot sole and finally the toes. In addition support is
Principles of human locomotion: a review*
J.L. Pons, Member, IEEE, J.C. Moreno, D. Torricelli and J.S. Taylor