Integrative Model of Drosophila Flight

This paper presents a framework for simulating the flight dynamics and control strategies of the fruit fly Drosophila melanogaster. The framework consists of five main components: an articulated rigid-body simulation, a model of the aerodynamic forces and moments, a sensory systems model, a control model, and an environment model. In the rigid-body simulation the fly is represented by a system of three rigid bodies connected by a pair of actuated ball joints. At each instant of the simulation, the aerodynamic forces and moments acting on the wings and body of the fly are calculated using an empirically derived quasi-steady model. The pattern of wing kinematics is based on data captured from high-speed video sequences. The forces and moments produced by the wings are modulated by deforming the base wing kinematics along certain characteristic actuation modes. Models of the fly’s visual and mechanosensory systems are used to generate inputs to a controller that sets the magnitude of each actuation mode, thus modulating the forces produced by the wings. This simulation framework provides a quantitative test bed for examining the possible control strategies employed by flying insects. Examples demonstrating pitch rate, velocity, altitude, and flight speed control, as well as visually guided centering in a corridor are presented.

[1]  Yuan-Cheng Fung,et al.  An introduction to the theory of aeroelasticity , 1955 .

[2]  M Egelhaaf,et al.  On the Computations Analyzing Natural Optic Flow: Quantitative Model Analysis of the Blowfly Motion Vision Pathway , 2005, The Journal of Neuroscience.

[3]  Erich Buchner,et al.  Behavioural Analysis of Spatial Vision in Insects , 1984 .

[4]  Patrick A. Shoemaker,et al.  Velocity constancy and models for wide-field visual motion detection in insects , 2005, Biological Cybernetics.

[5]  G K Taylor,et al.  Mechanics and aerodynamics of insect flight control , 2001, Biological reviews of the Cambridge Philosophical Society.

[6]  J. P. Lindemann,et al.  Function of a Fly Motion-Sensitive Neuron Matches Eye Movements during Free Flight , 2005, PLoS biology.

[7]  Martin Egelhaaf,et al.  Visual afferences to flight steering muscles controlling optomotor responses of the fly , 1989, Journal of Comparative Physiology A.

[8]  M. Dickinson,et al.  Summation of visual and mechanosensory feedback in Drosophila flight control , 2004, Journal of Experimental Biology.

[9]  Titus R. Neumann Modeling Insect Compound Eyes: Space-Variant Spherical Vision , 2002, Biologically Motivated Computer Vision.

[10]  M. Dickinson,et al.  Wing rotation and the aerodynamic basis of insect flight. , 1999, Science.

[11]  M. Dickinson,et al.  Spanwise flow and the attachment of the leading-edge vortex on insect wings , 2001, Nature.

[12]  Alexander Borst,et al.  Principles of visual motion detection , 1989, Trends in Neurosciences.

[13]  U. Grünert,et al.  Campaniform sensilla of Calliphora vicina (Insecta, Diptera) , 2004, Zoomorphology.

[14]  G. Nalbach The halteres of the blowfly Calliphora , 1993, Journal of Comparative Physiology A.

[15]  R. Durikovic,et al.  Human hand model based on rigid body dynamics , 2004 .

[16]  T. Collett,et al.  Visual control of flight behaviour in the hoverflySyritta pipiens L. , 1975, Journal of comparative physiology.

[17]  M. Dickinson,et al.  The control of flight force by a flapping wing: lift and drag production. , 2001, The Journal of experimental biology.

[18]  A. Straw,et al.  A `bright zone' in male hoverfly (Eristalis tenax) eyes and associated faster motion detection and increased contrast sensitivity , 2006, Journal of Experimental Biology.

[19]  Roger C. Hardie,et al.  Light Adaptation in Drosophila Photoreceptors , 2001, The Journal of general physiology.

[20]  C. Ellington The Aerodynamics of Hovering Insect Flight. II. Morphological Parameters , 1984 .

[21]  Dario Floreano,et al.  Vision-based Altitude and Pitch Estimation for Ultra-light Indoor Aircraft , 2006 .

[22]  Brett Browning,et al.  Accurate and flexible simulation for dynamic, vision-centric robots , 2004, Proceedings of the Third International Joint Conference on Autonomous Agents and Multiagent Systems, 2004. AAMAS 2004..

[23]  Brian Mirtich,et al.  Fast and Accurate Computation of Polyhedral Mass Properties , 1996, J. Graphics, GPU, & Game Tools.

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

[25]  R.M. Murray,et al.  Vision as a compensatory mechanism for disturbance rejection in upwind flight , 2004, Proceedings of the 2004 American Control Conference.

[26]  Mandyam V. Srinivasan,et al.  Motion detection in insect orientation and navigation , 1999, Vision Research.

[27]  M. Dickinson,et al.  The aerodynamic effects of wing rotation and a revised quasi-steady model of flapping flight. , 2002, The Journal of experimental biology.

[28]  C. Ellington THE AERODYNAMICS OF HOVERING INSECT FLIGHT. V. A VORTEX THEORY , 1984 .

[29]  M. Dickinson,et al.  The effect of advance ratio on the aerodynamics of revolving wings , 2004, Journal of Experimental Biology.

[30]  M. Dickinson,et al.  The correlation between wing kinematics and steering muscle activity in the blowfly Calliphora vicina. , 2001, The Journal of experimental biology.

[31]  Charles M Higgins,et al.  The computational basis of an identified neuronal circuit for elementary motion detection in dipterous insects. , 2004, Visual neuroscience.

[32]  Stéphane Viollet,et al.  Toward Optic Flow Regulation for Wall-Following and Centring Behaviours , 2006 .

[33]  P. Libby,et al.  Two-dimensional Problems in Hydrodynamics and Aerodynamics , 1965 .

[34]  M. Dickinson,et al.  Haltere Afferents Provide Direct, Electrotonic Input to a Steering Motor Neuron in the Blowfly, Calliphora , 1996, The Journal of Neuroscience.

[35]  C. Ellington The Aerodynamics of Hovering Insect Flight. VI. Lift and Power Requirements , 1984 .

[36]  M. Dickinson,et al.  The influence of wing–wake interactions on the production of aerodynamic forces in flapping flight , 2003, Journal of Experimental Biology.

[37]  B. Hassenstein,et al.  Systemtheoretische Analyse der Zeit-, Reihenfolgen- und Vorzeichenauswertung bei der Bewegungsperzeption des Rüsselkäfers Chlorophanus , 1956 .

[38]  N. Strausfeld Atlas of an Insect Brain , 1976, Springer Berlin Heidelberg.

[39]  James Sean Humbert,et al.  Bio-inspired visuomotor convergence in navigation and flight control systems , 2005 .

[40]  C. Ellington The Aerodynamics of Hovering Insect Flight. IV. Aeorodynamic Mechanisms , 1984 .

[41]  M. Dickinson,et al.  UNSTEADY AERODYNAMIC PERFORMANCE OF MODEL WINGS AT LOW REYNOLDS NUMBERS , 1993 .

[42]  A. Dubs,et al.  The dynamics of phototransduction in insects , 1984, Journal of Comparative Physiology A.

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

[44]  C. Ellington The Aerodynamics of Hovering Insect Flight. III. Kinematics , 1984 .

[45]  S. Shankar Sastry,et al.  Flapping flight for biomimetic robotic insects: part I-system modeling , 2006, IEEE Transactions on Robotics.

[46]  S. Shankar Sastry,et al.  Flapping flight for biomimetic robotic insects: part II-flight control design , 2006, IEEE Transactions on Robotics.

[47]  J. Pringle The gyroscopic mechanism of the halteres of Diptera , 1948, Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences.

[48]  M. Coutinho Dynamic Simulations of Multibody Systems , 2001, Springer New York.

[49]  D. Stewart,et al.  AN IMPLICIT TIME-STEPPING SCHEME FOR RIGID BODY DYNAMICS WITH INELASTIC COLLISIONS AND COULOMB FRICTION , 1996 .

[50]  C. David The relationship between body angle and flight speed in free‐flying Drosophila , 1978 .

[51]  Michael H Dickinson,et al.  The Initiation and Control of Rapid Flight Maneuvers in Fruit Flies1 , 2005, Integrative and comparative biology.

[52]  R. Hengstenberg Mechanosensory control of compensatory head roll during flight in the blowflyCalliphora erythrocephala Meig. , 1988, Journal of Comparative Physiology A.

[53]  A. Snyder Physics of Vision in Compound Eyes , 1979 .

[54]  Dickinson,et al.  THE EFFECTS OF WING ROTATION ON UNSTEADY AERODYNAMIC PERFORMANCE AT LOW REYNOLDS NUMBERS , 1994, The Journal of experimental biology.

[55]  M. Dickinson,et al.  Haltere-mediated equilibrium reflexes of the fruit fly, Drosophila melanogaster. , 1999, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[56]  S. N. Fry,et al.  The Aerodynamics of Free-Flight Maneuvers in Drosophila , 2003, Science.

[57]  R. Hengstenberg,et al.  Estimation of self-motion by optic flow processing in single visual interneurons , 1996, Nature.

[58]  M. Dickinson,et al.  Force production and flow structure of the leading edge vortex on flapping wings at high and low Reynolds numbers , 2004, Journal of Experimental Biology.

[59]  Victor B. Zordan,et al.  Physically based grasping control from example , 2005, SCA '05.

[60]  Roger C. Hardie,et al.  Light Adaptation in Drosophila Photoreceptors: I. Response Dynamics and Signaling Efficiency at 25°C , 2001 .