Cosine tuning determines plantarflexors' activities during human upright standing and is affected by incomplete spinal cord injury.

Plantarflexors such as the soleus (SOL) and medial gastrocnemius (MG) play key roles in controlling bipedal stance; however, how the central nervous system controls the activation levels of these plantarflexors is not well understood. Here we investigated how the central nervous system controls the plantarflexors' activation level during quiet standing in a cosine tuning manner where the maximal activation is achieved in a preferred direction (PD). Furthermore, we investigated how spinal cord injury affects these plantarflexors' activations. Thirteen healthy adults (AB) and thirteen individuals with chronic, incomplete spinal cord injury (iSCI) performed quiet standing trials. Their body kinematics and kinetics as well as electromyography signals from the MG and SOL were recorded. In the AB group, we found that the plantarflexors followed the cosine tuning manner during quiet standing. That is, MG was most active when the ratio of plantarflexion torque to knee extension torque was ~2:-3, whereas SOL was most active when the ratio was ~2:1. This suggests that the SOL muscle, despite being a monoarticular muscle, is sensitive to both ankle plantarflexion and knee extension during quiet standing. The difference in the PDs accounts for the phasic activity of MG and for the tonic activity of SOL. Unlike the AB group, the MG's activity was similar to the SOL's activity in the iSCI group, and the SOL PDs were similar to those in the AB group. This result suggests that chronic iSCI affects the control strategy, i.e., cosine tuning, for MG, which may affect standing balance in individuals with iSCI. Soleus muscle shows a tonic activity whereas medial gastrocnemius muscle shows a phasic activity during quiet standing. Cosine tuning and their preferred direction account for the different muscle activation patterns between these two muscles. In individuals with chronic incomplete spinal cord injury, the preferred direction of gastrocnemius medial head is affected, which may result in their deteriorated standing balance.