Locomotor strategy for pedaling: muscle groups and biomechanical functions

A group of coexcited muscles alternating with another group is a common element of motor control, including locomotor pattern generation. This study used computer simulation to investigate human pedaling with each muscle assigned at times to a group. Simulations were generated by applying patterns of muscle excitations to a musculoskeletal model that includes the dynamic properties of the muscles, the limb segments, and the crank load. Raasch et al. showed that electromyograms, pedal reaction forces, and limb and crank kinematics recorded during maximum-speed start-up pedaling could be replicated with two signals controlling the excitation of four muscle groups (1 group alternating with another to form a pair). Here a four-muscle-group control also is shown to replicate steady pedaling. However, simulations show that three signals controlling six muscle groups (i.e., 3 pairs) is much more biomechanically robust, such that a wide variety of forward and backward pedaling tasks can be executed well. We found the biomechanical functions necessary for pedaling, and how these functions can be executed by the muscle groups. Specifically, the phasing of two pairs with respect to limb extension and flexion and the transitions between extension and flexion do not change with pedaling direction. One pair of groups (uniarticular hip and knee extensors alternating with their anatomic antagonists) generates the energy required for limb and crank propulsion during limb extension and flexion, respectively. In the second pair, the ankle plantarflexors transfer the energy from the limb inertia to the crank during the latter part of limb extension and the subsequent limb extension-to-flexion transition. The dorsiflexors alternate with the plantarflexors. The phasing of the third pair (the biarticular thigh muscles) reverses with pedaling direction. In forward pedaling, the hamstring is excited during the extension-to-flexion transition and in backward pedaling during the opposite transition. In both cases hamstrings propel the crank posteriorly through the transition. Rectus femoris alternates with hamstrings and propels the crank anteriorly through the transitions. With three control signals, one for each pair of groups, different cadences (or power outputs) can be achieved by adjusting the overall excitatory drive to the pattern generating elements, and different pedaling goals (e.g., smooth, or energy-efficient pedaling; 1- or 2-legged pedaling) by adjusting the relative excitation levels among the muscle groups. These six muscle groups are suggested to be elements of a general strategy for pedaling control, which may be generally applicable to other human locomotor tasks.