Turning manoeuvres in free‐flying locusts: Two‐channel radio‐telemetric transmission of muscle activity

Kutsch, W.; Berger, Sebastian; Kautz, H.

doi:10.1002/jez.a.10297

Cited by 16 publications

(16 citation statements)

References 52 publications

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“…Until now, the wireless transmission of muscle potentials has been applied for freely flying desert locusts (Fischer and Kutsch, 1999;Kutsch, 2002;Kutsch et al, 2003) and hawkmoths (Ando and Kanzaki, 2004). The flight muscle activities and wing kinematics [such as wingbeat frequency, the firing relationship of elevation muscles and the correlation between muscle activities and body/wing kinematics (Kutsch, 2002;Kutsch et al, 2003)] during free flight of locusts could be different from those in tethered flight. As for hawkmoths during free flight, the flight muscle activities are much less changeable than those during tethered flight, suggesting that slight modulation of the motor output pattern and the subsequent wing kinematics might be adequate for free flight manoeuvres.…”

Section: Introductionmentioning

confidence: 99%

Active control of free flight manoeuvres in a hawkmoth, Agrius convolvuli

Wang

Ando

Kanzaki

2008

Journal of Experimental Biology

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SUMMARYBy combining optical triangulation with the comb-fringe technique and dual-channel telemetry, wing kinematics and body attitudes accompanying muscle activities of free-flying male hawkmoths were recorded synchronously when they performed flight manoeuvres elicited by a female sex pheromone. The results indicate that the wing leading edge angular position at the ventral stroke reversal, which can be decomposed by two orthogonal angular parameters (a flapping angle and a deviation angle), is well controllable. Two specific flight muscles, the dorsal-ventral muscle (DVM, indirect muscle, a wing elevator) and the third axillary muscle (3AXM, direct muscle, a wing retractor), can modulate the flapping angle and the deviation angle, respectively, by means of regulating the firing timing of muscle activities. The firing timing can be expressed by the firing latency absolutely, which is just before the timing of ventral stroke reversal. The results illustrate that lengthening the firing latency of the DVM and of the 3AXM can increase the flapping angle and the deviation angle, respectively, which both strengthen the downstroke at the ventral stroke reversal. The relationship of bilateral asymmetry shows that the bilateral differences in the firing latency of the DVM and of the 3AXM will cause bilateral differences in the wing position, which accompany the variations of yaw and roll angles in time course. This implies the contribution of the two muscles to active steering controls during turning or banking, though the DVM being an indirect muscle was generally treated as a power generator. Finally, the relationship between the pitch angle and the 3AXM latency, deduced from the relationships between the pitch angle and the deviation angle and between the deviation angle and the 3AXM latency, shows that lengthening the 3AXM latency can increase the pitch angle at the ventral stroke reversal by moving the wing tip far away from the centre of gravity of the body, which indicates a functional role of the 3AXM in active pitching control.

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Section: Introductionmentioning

confidence: 99%

Active control of free flight manoeuvres in a hawkmoth, Agrius convolvuli

Wang

Ando

Kanzaki

2008

Journal of Experimental Biology

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“…Whereas a dynamics approach directly links the motions of a system with the forces that produce them, a kinematics approach treats motions in isolation from the forces that produce them. This means that a kinematics approach is necessarily correlative: we know how body kinematics correlate with wing kinematics (Wakeling & Ellington 1997), we know how aerodynamic force production correlates with wing kinematics (Lehmann & Dickinson 1998) and we even know how wing and body kinematics are correlated with the firing of individual flight muscles in free flight (Kutsch et al 2003)-but our understanding of the motions nevertheless rests at the level of statistical correlation, rather than at the level of a physical model of the underlying mechanics. By providing this functional link between forces and motions, a flight dynamics approach provides a new, physically proper, paradigm for understanding insect flight control.…”

Section: Introductionmentioning

confidence: 99%

Nonlinear time-periodic models of the longitudinal flight dynamics of desert locustsSchistocerca gregaria

Taylor

Żbikowski

2005

J. R. Soc. Interface.

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Previous studies of insect flight control have been statistical in approach, simply correlating wing kinematics with body kinematics or force production. Kinematics and forces are linked by Newtonian mechanics, so adopting a dynamics-based approach is necessary if we are to place the study of insect flight on its proper physical footing. Here we develop semi-empirical models of the longitudinal flight dynamics of desert locusts Schistocerca gregaria. We use instantaneous force-moment measurements from individual locusts to parametrize the nonlinear rigid body equations of motion. Since the instantaneous forces are approximately periodic, we represent them using Fourier series, which are embedded in the equations of motion to give a nonlinear time-periodic (NLTP) model. This is a proper mathematical generalization of an earlier linear-time invariant (LTI) model of locust flight dynamics, developed using previously published time-averaged versions of the instantaneous force recordings. We perform various numerical simulations, within the fitted range of the model, and across the range of body angles used by free-flying locusts, to explore the likely behaviour of the locusts upon release from the tether. Solutions of the NLTP models are compared with solutions of the nonlinear time-invariant (NLTI) models to which they reduce when the periodic terms are dropped. Both sets of models are unstable and therefore fail to explain locust flight stability fully. Nevertheless, whereas the measured forces include statistically significant harmonic content up to about the eighth harmonic, the simulated flight trajectories display no harmonic content above the fundamental forcing frequency. Hence, manoeuvre control in locusts will not directly reflect subtle changes in the higher harmonics of the wing beat, but must operate on a coarser time-scale. A state-space analysis of the NLTP models reveals orbital trajectories that are impossible to capture in the LTI and NLTI models, and inspires the hypothesis that asymptotic orbital stability is the proper definition of stability in flapping flight. Manoeuvre control on the scale of more than one wing beat would then consist in exciting transients from one asymptotically stable orbit to another. We summarize these hypotheses by proposing a limit-cycle analogy for flapping flight control and suggest experiments for verification of the limit-cycle control analogy hypothesis.

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“…However, in contrast to most previous studies that categorized wingbeat kinematics (for review, see Taylor, 2001), as well as muscle activity (Kutsch et al, 2003;Spüler and Heide, 1978;Thüring, 1986;Waldman and Zarnack, 1988), according to the amount of force or torque produced, we organized our analysis based on particular features of wing motion we previously correlated with patterns of steering muscle activity. These features were 'downstroke deviation', a correlate of basalare muscle activity, and 'mode', a correlate of activity in the pteralae III and pteralae I muscles (Balint and Dickinson, 2001).…”

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confidence: 99%

Neuromuscular control of aerodynamic forces and moments in the blowfly,Calliphora vicina

Balint

Dickinson

2004

Journal of Experimental Biology

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SUMMARY Flies are among the most agile of flying insects, a capacity that ultimately results from their nervous system's control over steering muscles and aerodynamic forces during flight. In order to investigate the relationships among neuromuscular control, musculo-skeletal mechanics and flight forces, we captured high-speed, three-dimensional wing kinematics of the blowfly, Calliphora vicina, while simultaneously recording electromyogram signals from prominent steering muscles during visually induced turns. We used the quantified kinematics to calculate the translational and rotational components of aerodynamic forces and moments using a theoretical quasi-steady model of force generation, confirmed using a dynamically scaled mechanical model of a Calliphora wing. We identified three independently controlled features of the wingbeat trajectory –downstroke deviation, dorsal amplitude and mode. Modulation of each of these kinematic features corresponded to both activity in a distinct steering muscle group and a distinct manipulation of the aerodynamic force vector. This functional specificity resulted from the independent control of downstroke and upstroke forces rather than the independent control of separate aerodynamic mechanisms. The predicted contributions of each kinematic feature to body lift, thrust, roll, yaw and pitch are discussed.

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Turning manoeuvres in free‐flying locusts: Two‐channel radio‐telemetric transmission of muscle activity

Cited by 16 publications

References 52 publications

Active control of free flight manoeuvres in a hawkmoth, Agrius convolvuli

Active control of free flight manoeuvres in a hawkmoth, Agrius convolvuli

Nonlinear time-periodic models of the longitudinal flight dynamics of desert locustsSchistocerca gregaria

Neuromuscular control of aerodynamic forces and moments in the blowfly,Calliphora vicina

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