Enthalpy efficiency of the soleus muscle explains improvements in running economy

During human running, the soleus, as the main plantar flexor muscle, generates the majority of the mechanical work through active shortening. The fraction of chemical energy that is converted into muscular work (i.e. the enthalpy efficiency) depends on the muscle shortening velocity. Here, we investigated the soleus muscle fascicle behavior during running with respect to the enthalpy efficiency as a mechanism that could explain previously reported improvements in running economy after exercise-induced increases of plantar flexor strength and Achilles tendon stiffness. Healthy amateur runners were randomly assigned to a control (n=10) or intervention group (n=13), which performed a specific 14-week muscle-tendon training. Significant increases in plantar flexor maximum strength (10%) and Achilles tendon stiffness (31%) yet reduced metabolic cost of running (4%) was found only in the intervention group (p<0.05). Following training, the soleus fascicle velocity profile throughout stance was altered, with the fascicles operating at a higher enthalpy efficiency during the phase of muscle-tendon unit lengthening (15%) and in average over stance (7%, p<0.05). These findings show that the improvements in energetic cost following increases in plantar flexor strength and Achilles tendon stiffness can be attributed to increased enthalpy efficiency of the operating soleus. This provides the first experimental evidence that the soleus enthalpy efficiency is a determinant of human running economy. Furthermore, the current results imply that the soleus energy production in the first part of the stance phase were the muscle-tendon unit is lengthening is crucial for the overall metabolic energy cost of running.