Falling with Style: Bats Perform Complex Aerial Rotations by Adjusting Wing Inertia

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.

[1]  Joseph W Bahlman,et al.  Glide performance and aerodynamics of non-equilibrium glides in northern flying squirrels (Glaucomys sabrinus) , 2013, Journal of The Royal Society Interface.

[2]  Berg,et al.  The moment of inertia of bird wings and the inertial power requirement for flapping flight , 1995, The Journal of experimental biology.

[3]  E. Revilla,et al.  A movement ecology paradigm for unifying organismal movement research , 2008, Proceedings of the National Academy of Sciences.

[4]  Ulla M. Norberg,et al.  Moments of Inertia of Bat Wings and Body , 1991 .

[5]  C. Ellington The Aerodynamics of Hovering Insect Flight. I. The Quasi-Steady Analysis , 1984 .

[6]  R. Norberg,et al.  The pterostigma of insect wings an inertial regulator of wing pitch , 1972, Journal of comparative physiology.

[7]  Z. J. Wang,et al.  Falling paper: Navier-Stokes solutions, model of fluid forces, and center of mass elevation. , 2004, Physical review letters.

[8]  James A Simmons,et al.  Interaction of vestibular, echolocation, and visual modalities guiding flight by the big brown bat, Eptesicus fuscus. , 2004, Journal of vestibular research : equilibrium & orientation.

[9]  K. Breuer,et al.  Upstroke wing flexion and the inertial cost of bat flight , 2012, Proceedings of the Royal Society B: Biological Sciences.

[10]  K. Breuer,et al.  Time-resolved wake structure and kinematics of bat flight , 2009 .

[11]  Jian Chen,et al.  Quantifying the complexity of bat wing kinematics. , 2008, Journal of theoretical biology.

[12]  David E. Orin,et al.  Robot dynamics: equations and algorithms , 2000, Proceedings 2000 ICRA. Millennium Conference. IEEE International Conference on Robotics and Automation. Symposia Proceedings (Cat. No.00CH37065).

[13]  D. Lentink,et al.  Vortex-wake interactions of a flapping foil that models animal swimming and flight , 2008, Journal of Experimental Biology.

[14]  Physiology Morphology Vertebrate Flight Mechanics Physiology Morphology Ecology And Evolution , 2016 .

[15]  S. N. Fry,et al.  The aerodynamics of hovering flight in Drosophila , 2005, Journal of Experimental Biology.

[16]  A. Hedenström,et al.  Bat flight: aerodynamics, kinematics and flight morphology , 2015, The Journal of Experimental Biology.

[17]  G. R. Spedding,et al.  The implications of low-speed fixed-wing aerofoil measurements on the analysis and performance of flapping bird wings , 2008, Journal of Experimental Biology.

[18]  T. Kane,et al.  A dynamical explanation of the falling cat phenomenon , 1969 .

[19]  Ara Arabyan,et al.  A distributed control model for the air-righting reflex of a cat , 1998, Biological Cybernetics.

[20]  Joseph W Bahlman,et al.  Bats go head-under-heels: the biomechanics of landing on a ceiling , 2009, Journal of Experimental Biology.

[21]  Charles P. Ellington,et al.  THE AERODYNAMICS OF HOVERING INSECT FLIGHT. , 2016 .

[22]  T. Hedrick,et al.  Wingbeat Time and the Scaling of Passive Rotational Damping in Flapping Flight , 2009, Science.

[23]  R. Full,et al.  Tail-assisted pitch control in lizards, robots and dinosaurs , 2012, Nature.

[24]  A. Biewener,et al.  Low speed maneuvering flight of the rose-breasted cockatoo (Eolophus roseicapillus). II. Inertial and aerodynamic reorientation , 2007, Journal of Experimental Biology.

[25]  Andrew A Biewener,et al.  Pigeons produce aerodynamic torques through changes in wing trajectory during low speed aerial turns , 2015, Journal of Experimental Biology.

[26]  Nachum Ulanovsky,et al.  Spatial cognition in bats and rats: from sensory acquisition to multiscale maps and navigation , 2015, Nature Reviews Neuroscience.

[27]  Sharon M Swartz,et al.  Kinematics of slow turn maneuvering in the fruit bat Cynopterus brachyotis , 2008, Journal of Experimental Biology.

[28]  Richard L Essner Three-dimensional launch kinematics in leaping, parachuting and gliding squirrels. , 2002, The Journal of experimental biology.

[29]  Sharon M Swartz,et al.  Whole-body kinematics of a fruit bat reveal the influence of wing inertia on body accelerations , 2011, Journal of Experimental Biology.

[30]  A. Hedenström,et al.  Leading-Edge Vortex Improves Lift in Slow-Flying Bats , 2008, Science.

[31]  Z. J. Wang,et al.  Fruit flies modulate passive wing pitching to generate in-flight turns. , 2009, Physical review letters.

[32]  V. M. Zat︠s︡iorskiĭ Kinematics of human motion , 1998 .

[33]  A Jusufi,et al.  Righting and turning in mid-air using appendage inertia: reptile tails, analytical models and bio-inspired robots , 2010, Bioinspiration & biomimetics.

[34]  J. W. Humberston Classical mechanics , 1980, Nature.

[35]  J. Hill,et al.  Bats: A Natural History , 1984 .

[36]  Donald C. Dunbar,et al.  Aerial maneuvers of leaping lemurs: The physics of whole‐body rotations while airborne , 1988, American journal of primatology.

[37]  R J Full,et al.  How animals move: an integrative view. , 2000, Science.

[38]  Gabriel Taubin,et al.  3D reconstruction of bat flight kinematics from sparse multiple views , 2011, 2011 IEEE International Conference on Computer Vision Workshops (ICCV Workshops).