Published: November 16, 2015 DOI: 10.1371/journal.pbio.1002297
The remarkable maneuverability of flying animals results from precise movements of their highly specialized wings. Bats have evolved an impressive capacity to control their flight, in large part due to their ability to modulate wing shape, area, and angle of attack through many independently controlled joints. Bat wings, however, also contain many bones and relatively large muscles, and thus the ratio of bats’ wing mass to their body mass is larger than it is for all other extant flyers. Although the inertia in bat wings would typically be associated with decreased aerial maneuverability, we show that bat maneuvers challenge this notion. We use a model-based tracking algorithm to measure the wing and body kinematics of bats performing complex aerial rotations. Using a minimal model of a bat with only six degrees of kinematic freedom, we show that bats can perform body rolls by selectively retracting one wing during the flapping cycle. We also show that this maneuver does not rely on aerodynamic forces, and furthermore that a fruit fly, with nearly massless wings, would not exhibit this effect. Similar results are shown for a pitching maneuver. Finally, we combine high-resolution kinematics of wing and body movements during landing and falling maneuvers with a 52-degree-of-freedom dynamical model of a bat to show that modulation of wing inertia plays the dominant role in reorienting the bat during landing and falling maneuvers, with minimal contribution from aerodynamic forces. Bats can, therefore, use their wings as multifunctional organs, capable of sophisticated aerodynamic and inertial dynamics not previously observed in other flying animals. This may also have implications for the control of aerial robotic vehicles.
Bats demonstrate remarkable agility in flight, reorienting from a horizontal flying position to a heels-over-head roosting position and recovering from aerial stumbles with ease. In this paper, we demonstrate that bats are able to execute these elegant maneuvers using primarily inertial forces, by controlled articulation of their heavy wings. Video from multiple high-speed cameras is used to create a digital representation of several flights in which bats either land on the ceiling or attempt to land and then recover from the subsequent fall. We observe that even at low speeds, when aerodynamic forces are likely to be small, the bats are able to reorient using combinations of asymmetric wing motions. A “minimal model” of a bat is proposed, which consists of a body and wings able to execute simplified motions. The dynamics of this minimal bat are then simulated and we are able to show that both roll and pitching motions, similar to those observed in the videos, can be achieved using specific sequences of wing motions. To confirm the general predictions of the minimal model, we simulate a fully articulated wing and body model, using as input the complex wing articulations measured from a series of 12 flights. We compare the simulated body orientation with the measured body orientation, finding excellent agreement between the two and thus supporting the hypothesis that inertial forces and not aerodynamic forces are largely responsible for these low-speed aerial maneuvers.