Collision avoidance of flying locusts: steering torques and behaviour

Summary 1. Obstacles approaching in the flight path trigger postural and wing kinematic adjustments in tethered flying locusts. We sought to confirm that these behaviours were steering behaviours by measuring the changes in the flight forces associated with their execution. We also investigated the coordination of these behaviours in the execution of collision avoidance manoeuvres and the effect of speed or size of the obstacle on the timing and magnitude of the response. 2. Locusts were tethered and suspended in a wind tunnel from orthogonally arranged leaf springs mounted with strain gauges. Lift and yaw torque could be monitored unambiguously. We also monitored a forward translation force which combined pitch and thrust. During flight, the locusts were videotaped from behind while targets of different sizes (5cm35 cm, 7cm37 cm, 9cm39cm, 11cm311cm) were transported towards the head at different speeds (1, 2, 3 or 4 ms21). 3. Angular asymmetry of the forewings during the downstroke with the right forewing high, and abdomen and hindleg movement to the left, were temporally associated with an increase in yaw torque to the left. With the left forewing high, abdomen and hindleg movement to the right were temporally associated with a decrease in yaw torque to the left. Obstacle avoidance behaviours could be associated with either an increase or a decrease in the pitch/thrust component. 4. Leg, abdomen and wingbeat alterations in response to the approach of an obstacle were independent but tightly coordinated. Slower approaches increased the magnitude of the responses. However, the size of the obstacle had no effect on the magnitude of the response. Slower and larger targets generated earlier reactions (i.e. locusts reacted when the targets were further from the head). 5. We conclude that the behaviours we have described were steering behaviours which would have directed the animal around an obstacle in its flight path, and that there were at least two strategies for collision avoidance associated with slowing or speeding flight. Leg, abdomen and wingbeat alterations formed a coherent avoidance response, the magnitude of which was dependent upon the time available for it to develop. We further conclude that the manoeuvre was not initiated at a constant time to collision and we propose that the avoidance strategy was to initiate the manoeuvre when the targets subtended more than 10˚ in the insect’s field of view.

[1]  M. Jensen Biology and physics of locust flight. III. The aerodynamics of locust flight , 1956, Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences.

[2]  K. D. Roeder The behaviour of free flying moths in the presence of artificial ultrasonic pulses , 1962 .

[3]  Boris Uvarov,et al.  Grasshoppers and Locusts: A Handbook of General Acridology. Vol. 1. Anatomy, Physiology, Phase Polymorphism, Introduction to Taxonomy. , 1967 .

[4]  J. Dugard Directional change in flying locusts , 1967 .

[5]  G. Varley,et al.  Grasshoppers and Locusts , 1967 .

[6]  J. Camhi Sensory Control of Abdomen Posture in Flying Locusts , 1970 .

[7]  Z. Waloff Orientation of flying locusts, Schistocerca gregaria (Forsk.), in migrating swarms , 1972 .

[8]  G. Pollack,et al.  Steering responses of flying crickets to sound and ultrasound: Mate attraction and predator avoidance. , 1978, Proceedings of the National Academy of Sciences of the United States of America.

[9]  M. Gewecke,et al.  Control of the horizontal flight‐course by air‐current sense organs in Locusta migratoria , 1978 .

[10]  M. Cloupeau,et al.  Direct Measurements of Instantaneous Lift in Desert Locust; Comparison with Jensen'S Experiments on Detached Wings , 1979 .

[11]  Lee Dn,et al.  The optic flow field: the foundation of vision. , 1980 .

[12]  C. Taylor Contribution of Compound Eyes and Ocelli to Steering Of Locusts in Flight: I. Behavioural Analysis , 1981 .

[13]  David N. Lee,et al.  Plummeting gannets: a paradigm of ecological optics , 1981, Nature.

[14]  J. Blondeau Aerodynamic Capabilities of Flies, as Revealed by a New Technique , 1981 .

[15]  H. Wagner Flow-field variables trigger landing in flies , 1982, Nature.

[16]  B. Mhl The role of proprioception in locust flight control: I. Asymmetry and coupling within the time pattern of motor units , 1985 .

[17]  D. E. Alexander Wind tunnel studies of turns by flying dragonflies. , 1986, The Journal of experimental biology.

[18]  D. Robert THE AUDITORY BEHAVIOUR OF FLYING LOCUSTS , 1989 .

[19]  M. Wortmann,et al.  ON THE SO-CALLED CONSTANT-LIFT REACTION OF MIGRATORY LOCUSTS , 1989 .

[20]  W. Rice ANALYZING TABLES OF STATISTICAL TESTS , 1989, Evolution; international journal of organic evolution.

[21]  R. Hoy,et al.  Ultrasound-induced yaw movements in the flying Australian field cricket (Teleogryllus oceanicus). , 1990, The Journal of experimental biology.

[22]  D D Yager,et al.  Ultrasound-triggered, flight-gated evasive maneuvers in the praying mantis Parasphendale agrionina. II. Tethered flight. , 1990, The Journal of experimental biology.

[23]  A. Baader THE POSTURE OF THE ABDOMEN DURING LOCUST FLIGHT: REGULATION BY STEERING AND VENTILATORY INTERNEURONES , 1990 .

[24]  R. Hoy,et al.  Leg-induced Steering in Flying Crickets , 1990 .

[25]  M. Fenton,et al.  Ultrasound-triggered, flight-gated evasive maneuvers in the praying mantis Parasphendale agrionina. I. Free flight. , 1990, The Journal of experimental biology.

[26]  D. Robert,et al.  I. Head movements and the organization of correctional manoeuvres , 1992 .

[27]  Daniel Robert,et al.  II. Acoustic avoidance manoeuvres and associated head movements, compared with correctional steering , 1992 .

[28]  D. N. Reye,et al.  WING MOVEMENTS ASSOCIATED WITH COLLISIONAVOIDANCE MANOEUVRES DURING FLIGHT IN THE LOCUST LOCUSTA MIGRATORIA , 1992 .

[29]  R. Hoy,et al.  Multisegmental analyses of acoustic startle in the flying cricket (Teleogryllus oceanicus): kinematics and electromyography. , 1992, The Journal of experimental biology.