Avoiding obstacles while intercepting a moving target: a miniature fly's solution.

The miniature robber fly Holcocephala fusca intercepts its targets with behaviour that is approximated by the proportional navigation guidance law. During predatory trials, we challenged the interception of H. fusca performance by placing a large object in its potential flight path. In response, H. fusca deviated from the path predicted by pure proportional navigation, but in many cases still eventually contacted the target. We show that such flight deviations can be explained as the output of two competing navigational systems: pure-proportional navigation and a simple obstacle avoidance algorithm. Obstacle avoidance by H. fusca is here described by a simple feedback loop that uses the visual expansion of the approaching obstacle to mediate the magnitude of the turning-away response. We name the integration of this steering law with proportional navigation 'combined guidance'. The results demonstrate that predatory intent does not operate a monopoly on the fly's steering when attacking a target, and that simple guidance combinations can explain obstacle avoidance during interceptive tasks.

Interception by two predatory fly species is explained by a proportional navigation feedback controller.

When aiming to capture a fast-moving target, animals can follow it until they catch up, or try to intercept it. In principle, interception is the more complicated strategy, but also more energy efficient. To study whether simple feedback controllers can explain interception behaviours by animals with miniature brains, we have reconstructed and studied the predatory flights of the robber fly Holcocephala fusca and killer fly Coenosia attenuata Although both species catch other aerial arthropods out of the air, Holcocephala contrasts prey against the open sky, while Coenosia hunts against clutter and at much closer range. Thus, their solutions to this target catching task may differ significantly. We reconstructed in three dimensions the flight trajectories of these two species and those of the presented targets they were attempting to intercept. We then tested their recorded performances against simulations. We found that both species intercept targets on near time-optimal courses. To investigate the guidance laws that could underlie this behaviour, we tested three alternative control systems (pure pursuit, deviated pursuit and proportional navigation). Only proportional navigation explains the timing and magnitude of fly steering responses, but with differing gain constants and delays for each fly species. Holcocephala uses a dimensionless navigational constant of N ≈ 3 with a time delay of ≈28 ms to intercept targets over a comparatively long range. This constant is optimal, as it minimizes the control effort required to hit the target. In contrast, Coenosia uses a constant of N ≈ 1.5 with a time delay of ≈18 ms, this setting may allow Coenosia to cope with the extremely high line-of-sight rotation rates, which are due to close target proximity, and thus prevent overcompensation of steering. This is the first clear evidence of interception supported by proportional navigation in insects. This work also demonstrates how by setting different gains and delays, the same simple feedback controller can yield the necessary performance in two different environments.

Responses of the two-spotted oak buprestid, Agrilus biguttatus (Coleoptera: Buprestidae), to host tree volatiles.

BACKGROUND: Agrilus bigutattus (Fabricius) is a forest pest of increasing importance in the United Kingdom. The larvae damage weakened native oaks and are thought to contribute to premature tree death. Suspected links with acute oak decline (AOD) are not yet confirmed, but AOD-predisposed trees appear to become more susceptible to A. biguttatus attack. Thus, management may be necessary for control of this insect. To explore the possibility of monitoring beetle populations by baited traps, the host tree volatiles regulating A. biguttatus-oak interactions were studied. RESULTS: Biologically active volatile organic compounds in dynamic headspace extracts of oak foliage and bark were identified initially by coupled gas chromatography-electroantennography (GC-EAG) and GC-mass spectrometry (GC-MS), and the structures were confirmed by GC coinjection with authentic compounds. Of two synthetic blends of these compounds comprising the active leaf volatiles, the simpler one containing three components evoked strongly positive behavioural responses in four-arm olfactometer tests with virgin females and males, although fresh leaf material was more efficient than the blend. The other blend, comprising a five-component mixture made up of bark volatiles, proved to be as behaviourally active for gravid females as bark tissue. CONCLUSIONS: These initial results on A. biguttatus chemical ecology reveal aspects of the role of attractive tree volatiles in the host-finding of beetles and underpin the development of semiochemically based surveillance strategies for this forest insect.