In vivo muscle force and elastic energy storage during steady-speed hopping of tammar wallabies (Macropus eugenii)
In order to evaluate the role of elastic energy recovery in the hopping of macropodids, in vivo measurements of muscle­tendon forces using buckle force transducers attached to the tendons of the gastrocnemius (G), plantaris (PL) and flexor digitorum longus (FDL) of tammar wallabies were made as the animals hopped on a treadmill at speeds ranging from 2.1 to 6.3 m s-1. These muscles and tendons constitute the main structures that are most important in energy storage and recovery. Electromyographic recordings from the lateral gastrocnemius and plantaris muscles, together with high-speed films (200 frames s-1) and video (60 fields s-1), were also used to correlate muscle activation and kinematic patterns of limb movement with force development. On the basis of in situ calibrations of the buckle transducers, we found that muscle forces and elastic energy storage increased with increased hopping speed in all three muscle­tendon units. Elastic energy recovery reached a maximum of 25 % of metabolic energy expenditure at 6.3 m s-1 and is probably greater than this at higher speeds. Force sharing among the three muscles was consistently maintained over this range of speeds in terms of recruitment. Although forces and stresses were generally comparable within the gastrocnemius and plantaris muscles, maximal tendon stresses were considerably greater in the gastrocnemius, because of its smaller cross-sectional area (peak muscle stress: 227 versus 262 kPa; peak tendon stress: 36 versus 32 MPa, G versus PL). As a result, energy storage was greatest in the gastrocnemius tendon despite its much shorter length, which limits its volume and, hence, energy storage capacity, compared with PL and FDL tendons. Forces and stresses (17 MPa maximum) developed within the FDL tendon were consistently much lower than those for the other two tendons. Peak stresses in these three tendons indicated safety factors of 3.0 for G, 3.3 for PL and 6.0 for FDL. The lower stresses developed within the tendons of the plantaris and, especially, the flexor digitorum longus may indicate the need to maintain sufficient stiffness for phalangeal control of foot placement, at the expense of reduced strain energy recovery.