SUMMARYFlight control in insects depends on self-induced image motion (optic flow), which the visual system must process to generate appropriate corrective steering maneuvers. Classic experiments in tethered insects applied rigorous system identification techniques for the analysis of turning reactions in the presence of rotating pattern stimuli delivered in open-loop. However, the functional relevance of these measurements for visual free-flight control remains equivocal due to the largely unknown effects of the highly constrained experimental conditions. To perform a systems analysis of the visual flight speed response under freeflight conditions, we implemented a 'one-parameter open-loop' paradigm using 'TrackFly' in a wind tunnel equipped with real-time tracking and virtual reality display technology. Upwind flying flies were stimulated with sine gratings of varying temporal and spatial frequencies, and the resulting speed responses were measured from the resulting flight speed reactions. To control flight speed, the visual system of the fruit fly extracts linear pattern velocity robustly over a broad range of spatio-temporal frequencies. The speed signal is used for a proportional control of flight speed within locomotor limits. The extraction of pattern velocity over a broad spatio-temporal frequency range may require more sophisticated motion processing mechanisms than those identified in flies so far. In Drosophila, the neuromotor pathways underlying flight speed control may be suitably explored by applying advanced genetic techniques, for which our data can serve as a baseline. Finally, the high-level control principles identified in the fly can be meaningfully transferred into a robotic context, such as for the robust and efficient control of autonomous flying micro air vehicles. Supplementary material available online at

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Behavioural control in many animals involves complex mechanisms with intricate sensorymotor feedback loops. Modelling allows functional aspects to be captured without relying on a description of the underlying complex, and often unknown, mechanisms. A wide range of engineering techniques are available for modelling, but their ability to describe time-continuous processes is rarely exploited to describe sensory-motor control mechanisms in biological systems. We performed a system identification of visual flight speed control in the fruitfly Drosophila, based on an extensive dataset of open-loop responses previously measured under free flight conditions. We identified a second-order under-damped control model with just six free parameters that well describes both the transient and steady-state characteristics of the open-loop data. We then used the identified control model to predict flight speed responses after a visual perturbation under closed-loop conditions and validated the model with behavioural measurements performed in free-flying flies under the same closed-loop conditions. Our system identification of the fruitfly's flight speed response uncovers the high-level control strategy of a fundamental flight control reflex without depending on assumptions about the underlying physiological mechanisms. The results are relevant for future investigations of the underlying neuromotor processing mechanisms, as well as for the design of biomimetic robots, such as micro-air vehicles.