How cockatiels (Nymphicus hollandicus) modulate pectoralis power output across flight speeds

The avian pectoralis muscle must produce a varying mechanical power output to achieve flight across a range of speeds (1-13 m s^-1). We used the natural variation in the power requirements with flight speed to investigate the mechanisms employed by cockatiels (Nymphicus hollandicus) to modulate muscle power output. We found that pectoralis contractile function in cockatiels was generally conserved across speed and over a wide range of aerodynamic power requirements. Despite the 2-fold range of variation in muscle power output, many aspects of muscle performance varied little: duration of muscle shortening was invariant, and overall wingbeat frequency and muscle strain varied to a lesser degree (1.2-fold and 1.4-fold, respectively) than muscle power or work. Power output was primarily modulated by muscle force (accounting for 65% of the variation) rather than by muscle strain, cycle frequency or changes in the timing of force production relative to muscle strain. Strain rate and electromyogram (EMG) results suggest that the additional force was provided via increasing pectoralis recruitment. Due to their effect on the transformation of muscle work into useful aerodynamic work, changes in wing position and orientation during the downstroke probably also affect the magnitude of muscle force developed for a given level of motor recruitment. Analysis of the variation in muscle force and airflow over the wing suggests that the coefficients of lift and drag of the wing vary 4-fold over the speed range examined in this study.