Where I have been explaining many of my previous measurements identifying flapping flight as aerodynamically inefficient by exploring the competing inertial costs of repeatedly accelerating wings with mass (see ‘flight’ section), this account does a poor job at accounting for the scaling of flapping gaits and flight strategies. An alternative account focussing on some simple costs and demands associated with activating muscle (see the ‘Big Idea’) does much better in accounting for the high-amplitude flaps of small birds, and the occurrence of bounding and flap-gliding flight strategies. The movie here takes about 15 minutes. The abstract below covers the main points, taken from the paper:
Usherwood, J.R. (2016). Physiological, aerodynamic and geometric constraints of flapping account for bird gaits, and bounding and flap-gliding flight strategies. J. Theor. Biol. doi: 10.1016/j.jtbi.2016.07.003
Aerodynamically economical flight is steady and level. The high-amplitude flapping and bounding flight style of many small birds departs considerably from any aerodynamic or purely mechanical optimum. Further, many large birds adopt a flap-glide flight style in cruising flight which is not consistent with purely aerodynamic economy. Here, an account is made for such strategies by noting a well-described, general, physiological cost parameter of muscle: the cost of activation. Small birds, with brief downstrokes, experience disproportionately high costs due to muscle activation for power during contraction as opposed to work. Bounding flight may be an adaptation to modulate mean aerodynamic force production in response to 1) physiological pressure to extend the duration of downstroke to reduce power demands during contraction; 2) the prevention of a low-speed downstroke due to the geometric constraints of producing thrust; 3) an aerodynamic cost to flapping with very low lift coefficients. In contrast, flap-gliding birds, which tend to be larger, adopt a strategy that reduces the physiological cost of work due both to activation and contraction efficiency. Flap-gliding allows, despite constraints to modulation of aerodynamic force lever-arm, 1) adoption of moderately large wing-stroke amplitudes to achieve suitable muscle strains, thereby reducing the activation costs for work; 2) reasonably quick downstrokes, enabling muscle contraction at efficient velocities, while being 3) prevented from very slow weight-supporting upstrokes due to the cost of performing ‘negative’ muscle work.