Measuring metabolic rate (MR) poses a formidable challenge in free-flying insects who cannot breathe into masks or be trained to fly in controlled settings. Consequently, flight MR has been predominantly measured on hovering or tethered insects flying in closed systems. Stable isotopes such as labelled water allow measurement of MR in free-flying animals but integrates the measurement over long periods exceeding the average flight duration of insects. Here, we applied the ‘bolus injection of isotopic 13 C Na-bicarbonate’ method to insects to measure their flight MR and report a 90% accuracy compared to respirometry. We applied the method on two beetle species, measuring MR during free flight and tethered flight in a wind tunnel. We also demonstrate the ability to repeatedly use the technique on the same individual. Therefore, the method provides a simple, reliable and accurate tool that solves a long-lasting limitation on insect flight research by enabling the measurement of MR during free flight.

The need for long dispersal flights can drive selection for behavioral, physiological and biomechanical mechanisms to reduce the energy spent flying. However, some energy loss during the transfer of momentum from the wing to the fluid is inevitable, and inherent to the fluid-wing interaction. Here, we analyzed these losses during the forward flight of the mango stem borer (Batocera rufomaculata). This relatively large beetle can disperse substantial distances in search of new host trees, and laboratory experiments have demonstrated continuous tethered flights that can last for up to an hour. We flew the beetles tethered in a wind tunnel and used high-speed videography to estimate the aerodynamic power from their flapping kinematics and particle image velocimetry (PIV) to evaluate drag and kinetic energy from their unsteady wakes. To account for tethering effects, we measured the forces applied by the beetles on the tether arm holding them in place. The drag of the flying beetle over the flapping cycle, estimated from the flow fields in the unsteady wake, showed good agreement with direct measurement of mean horizontal force. Both measurements showed that total drag during flight is ∼5-fold higher than the parasite drag on the body. The aerodynamic power estimated from both the motion of the wings, using a quasi-steady blade-element model, and the kinetic energy in the wake, gave mean values of flight-muscle mass-specific power of 87 and 65 W kg muscle−1, respectively. A comparison of the two values suggests that ∼25% of the energy is lost within the fluid due to turbulence and heat. The muscle mass-specific power found here is low relative to the maximal power output reported for insect flight muscles. This can be attributed to reduced weight support during tethered flight or to operation at submaximal output that may ensure a supply of metabolic substrates to the flight muscles, thus delaying their fatigue during long-distance flights.

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The distance a small insect moves through the air during a jump is limited by the launch velocity at take-off and by air resistance. The launch velocity is limited by the length of the jumping legs and the maximum power that the jump apparatus can provide for pushing against the ground. The effect of air resistance is determined by the insect mass-to-area ratio. Both limitations are highly dependent on body size, making high jumps a challenge for smaller insects. We studied both effects in the tiny Encyrtid wasp Males are smaller than females (mean body length 1.2 and 1.8 mm, respectively), but both sexes take off in a powerful jump. Using high-speed cameras, we analyzed the relationship between take-off kinematics and distance traveled through the air. We show that the velocity, acceleration and mass-specific power when leaving the ground places among the most prominent jumpers of the insect world. However, the absolute distance moved through the air is modest compared with other jumping insects, as a result of air resistance acting on the small body. A biomechanical model suggests that air resistance reduces the jump distance of these insects by 49% compared with jumping in the absence of air resistance. The effect of air resistance is more pronounced in the smaller males, resulting in a segregation of the jumping performance between sexes. The limiting effect of air resistance is inversely proportional to body mass, seriously constraining jumping as a form of moving through the air in these and other small insects.

Rose chafer beetles (Protetia cuprea) are pollinators as well as agricultural pests, flying between flowers and trees while foraging for pollen and fruits. Calculating the energy they expend on flying during foraging activity faces the challenge of measuring the metabolic rate (MR) of free‐flying insects in an open space. We overcame this challenge by using the bolus injection of 13C Na‐bicarbonate technique to measure their metabolic energy expenditure while flying in a large flight arena. Concurrently, we tracked the insects with high‐speed cameras to extract their flight trajectory, from which we calculated the mechanical power invested in flying for each flight bout. We found that the chemical (metabolic) energy input converted to mechanical flight energy output at a mean efficiency of 10.4% ± 5.2%, with a trend of increased efficiency in larger conspecifics (efficiency scaled with body mass to the power of 1.4). The transition in the summer from a diet of pollen to that of fruits may affect the energy budget available for foraging. Starved P. cuprea, feeding on apples ad libitum, increased their body mass by an average of 6% in 2 h. According to our calculations, such a meal can power a 630‐m flight (assuming a carbohydrate assimilation efficiency of 90%). Pollen, with a low water and carbohydrate content but rich in proteins and lipids, has a higher caloric content and should assimilate differently when converting food to flight fuel. The high cost of aerial locomotion is inherent to the foraging behavior of rose chafers, explaining their short flight bouts followed by prolonged feeding activity.