Dynamics of quadrupedal locomotion of monkeys: implications for central control

Abstract

We characterized the three-dimensional kinematics and dynamics of quadrupedal gait of young adult rhesus and cynomolgus monkeys while they walked with diagonal and lateral gaits at 0.4–1.0 m/s on a treadmill. Rigid bodies on the wrist, ankle, and back were monitored by an optical motion detection system (Optotrak). Kinematic data could be normalized using characteristic stride length, reducing variance due to different gait styles, to emphasize common characteristics of swing and stance parameters among animals. Mean swing phase durations fell as walking speed increased, but the swing phase durations increased at each walking velocity as a linear function of increases in amplitude, thereby following a main sequence relationship. The phase plane trajectories of the swing phases, i.e., plots of the relation of the rising and falling limb velocity to limb position in the sagittal (X–Z) plane, had unique dynamic characteristics. Trajectories were separable at each walking velocity and increases in swing amplitude were linearly related to peak swing velocities, thus comprising main sequences. We infer that the swing phase dynamics are set by central neural mechanisms at the onset of the swing phases according to the intended amplitude, which in turn is based on the walking velocity in the stance phases. From the many dynamic similarities between swing phases and rapid eye movements, we further suggest that the swing phases may be generated by neural mechanisms similar to those that produce saccades and quick phases of nystagmus from a signal related to sensed or desired walking velocity.

Appendix

Kinematic data were normalized relative to the average stride length (characteristic length). This characteristic length was denoted as L _c , the stride frequency as F _stride, and the swing and stance durations as T _swing and T _stance, respectively. From this, with the acceleration of gravity g, the normalized stride length $L^{n}_{{{\text{stride}}}}$, stride frequency $F^{n}_{{{\text{stride}}}}$, and swing/stance durations ( $T^{n}_{{{\text{swing}}}}$ and $T^{n}_{{{\text{stance}}}}$) were represented as

$$ L^{n}_{{{\text{stride}}}} = \frac{{L_{{{\text{stride}}}} }} {{L_{c} }} $$

(1)

$$ F^{n}_{{{\text{stride}}}} = F_{{{\text{stride}}}} \cdot {\sqrt {\frac{{L_{c} }} {g}} } $$

(2)

$$ T^{n}_{{{\text{swing}}}} = T_{{{\text{swing}}}} \cdot {\sqrt {\frac{g} {{L_{c} }}} } $$

(3)

$$ T^{n}_{{{\text{stance}}}} = T_{{{\text{stance}}}} \cdot {\sqrt {\frac{g} {{L_{c} }}} } $$

(4)

Similarly the original walking speed, V, was converted into a normalized walking speed V ⁿ and represented as

$$ V^{n} = \frac{V} {{{\sqrt {g \cdot L_{c} } }}} $$

(5)