The mechanical power output of the flight muscles of blue-breasted quail (Coturnix chinensis) during take-off.

Blue-breasted quail (Coturnix chinensis) were filmed during take-off flights. By tracking the position of the centre of mass of the bird in three dimensions, we were able to calculate the power required to increase the potential and kinetic energy. In addition, high-speed video recordings of the position of the wings over the course of the wing stroke, and morphological measurements, allowed us to calculate the aerodynamic and inertial power requirements. The total power output required from the pectoralis muscle was, on average, 390 W kg(-1), which was similar to the highest measurements made on bundles of muscle fibres in vitro (433 W kg(-1)), although for one individual a power output of 530 W kg(-1) was calculated. The majority of the power was required to increase the potential energy of the body. The power output of these muscles is the highest yet found for any muscle in repetitive contractions. We also calculated the power requirements during take-off flights in four other species in the family Phasianidae. Power output was found to be independent of body mass in this family. However, the precise scaling of burst power output within this group must await a better assessment of whether similar levels of performance were measured across the group. We extended our analysis to one species of hawk, several species of hummingbird and two species of bee. Remarkably, we concluded that, over a broad range of body size (0.0002-5 kg) and contractile frequency (5-186 Hz), the myofibrillar power output of flight muscles during short maximal bursts is very high (360-460 W kg(-1)) and shows very little scaling with body mass. The approximate constancy of power output means that the work output varies inversely with wingbeat frequency and reaches values of approximately 30-60 J kg(-1) in the largest species.

The novel aerodynamics of insect flight: applications to micro-air vehicles.

The wing motion in free flight has been described for insects ranging from 1 to 100 mm in wingspan. To support the body weight, the wings typically produce 2-3 times more lift than can be accounted for by conventional aerodynamics. Some insects use the fling mechanism: the wings are clapped together and then flung open before the start of the downstroke, creating a lift-enhancing vortex around each wing. Most insects, however, rely on a leading-edge vortex (LEV) created by dynamic stall during flapping; a strong spanwise flow is also generated by the pressure gradients on the flapping wing, causing the LEV to spiral out to the wingtip. Technical applications of the fling are limited by the mechanical damage that accompanies repeated clapping of the wings, but the spiral LEV can be used to augment the lift production of propellers, rotors and micro-air vehicles (MAVs). Design characteristics of insect-based flying machines are presented, along with estimates of the mass supported, the mechanical power requirement and maximum flight speeds over a wide range of sizes and frequencies. To support a given mass, larger machines need less power, but smaller ones operating at higher frequencies will reach faster speeds.

The mechanics of flight in the hawkmoth Manduca sexta. I. Kinematics of hovering and forward flight.

High-speed videography was used to record sequences of individual hawkmoths in free flight over a range of speeds from hovering to 5 ms-1. At each speed, three successive wingbeats were subjected to a detailed analysis of the body and wingtip kinematics and of the associated time course of wing rotation. Results are presented for one male and two female moths. The clearest kinematic trends accompanying increases in forward speed were an increase in stroke plane angle and a decrease in body angle. The latter may have resulted from a slight dorsal shift in the area swept by the wings as the supination position became less ventral with increasing speed. These trends were most pronounced between hovering and 3 ms-1, and the changes were gradual; there was no distinct gait change of the kind observed in some vertebrate fliers. The wing rotated as two functional sections: the hindwing and the portion of the forewing with which it is in contact, and the distal half of the forewing. The latter displayed greater fluctuation in the angle of rotation, especially at the lower speeds. As forward speed increased, the discrepancy between the rotation angles of the two halfstrokes, and of the two wing sections, became smaller. The downstroke wing torsion was set early in the halfstroke and then held constant during the translational phase.

The mechanics of flight in the hawkmoth Manduca sexta. II. Aerodynamic consequences of kinematic and morphological variation.

Mean lift coefficients have been calculated for hawkmoth flight at a range of speeds in order to investigate the aerodynamic significance of the kinematic variation which accompanies changes in forward velocity. The coefficients exceed the maximum steady-state value of 0.71 at all except the very fastest speeds, peaking at 2.0 or greater between 1 and 2 ms-1. Unsteady high-lift mechanisms are therefore most important during hovering and slow forward flight. In combination with the wingtip paths relative to the surrounding air, the calculated mean lift coefficients illustrate how the relative contributions of the two halfstrokes to the force balance change with increasing forward speed. Angle of incidence data for fast forward flight suggest that the sense of the circulation is not reversed between the down- and upstrokes, indicating a flight mode qualitatively different from that proposed for lower-speed flight in the hawkmoth and other insects. The mid-downstroke angle of incidence is constant at 30-40 degrees across the speed range. The relationship between power requirements and flight speed is explored; above 5 ms-1, further increases in forward velocity are likely to be constrained by available mechanical power, although problems with thrust generation and flight stability may also be involved. Hawkmoth wing and body morphology, and the differences between males and females, are evaluated in aerodynamic terms. Steady-state force measurements show that the hawkmoth body is amongst the most streamlined for any insect.

Unsteady aerodynamics of insect flight.

Over the past decade, the importance of unsteady aerodynamic mechanisms for flapping insect flight has become widely recognised. Even at the fastest flight speeds, the old quasi-steady aerodynamic interpretation seems inadequate to explain the extra lift produced by the wings. Recent experiments on rigid model wings have confirmed the effectiveness of several postulated high-lift mechanisms. Delayed stall can produce extra lift for several chords of travel during the translational phases of the wingbeat. Lift can also be enhanced by circulation created during pronation and supination by rotational mechanisms: the fling/peel, the near fling/peel and isolated rotation. These studies have revealed large leading-edge vortices which contribute to the circulation around the wing, augmenting the lift. The mechanisms show distinctive patterns of vortex shedding from leading and trailing edges. The results of flow visualization experiments on tethered insects are reviewed in an attempt to identify the high-lift mechanisms actually employed. The fling/peel mechanism is clearly used by some insects. The near fling/peel is the wing motion most commonly observed, but evidence for the production of high lift remains indirect. For many insects, lift on the upstroke probably results from delayed stall instead of the flex mechanism of isolated rotation. The large leading-edge vortices from experiments on rigid model wings are greatly reduced or missing around the real insect wings, often making the identification of aerodynamic mechanisms inconclusive. A substantial spanwise flow component has been detected over the aerodynamic upper wing surface, which should transport leading-edge vorticity towards the wingtip before it has much time to roll up. This spanwise transport, arising from centrifugal acceleration, is probably a general phenomenon for flapping insect flight. It should reduce and stabilise any leading-edge vortices that are present, which is essential for preventing stall and maintaining the circulation of high-lift mechanisms during translation.

Beyond the vertebrates: achieving maximum power during flight in insects and hummingbirds.

Hummingbirds and insects are clearly extreme aerobic athletes. Hovering hummingbirds and insects exhibit the highest mass-specific metabolic rates found in vertebrates and invertebrates, respectively. Both groups of fliers have high mitochondrial volume densities in their locomotor muscles, but these do not exceed 35-40% of the fiber volume, presumably from a need to conserve myofibrils for force generation. A possible adaptation to this constraint is the observed greater packing of the inner mitochondrial membranes than occurs in mammalian mitochondria. Both hummingbirds and insects show higher rates of oxygen consumption per unit volume of mitochondria than do mammals. Additionally, volume-specific mitochondrial oxygen consumption in insects increases as body size decreases, unlike the size-independent pattern in mammals. Aerodynamic analysis of power output during hovering flight strongly suggests that both insects and hummingbirds operate with considerable elastic storage of kinetic energy, thereby decreasing their inertial power requirements. Both groups appear to hover with muscle power output close to 100 W kg-1. Muscle efficiency in hummingbirds is near 10%; in insects, muscle efficiency varies with body size, but at similarly low values. Scaling of efficiency with body size has also been reported in terrestrial mammals, suggesting a possible common mechanism. Both groups of hovering fliers can markedly increase their metabolic power inputs and mechanical power outputs above those required for basic hovering flight. These elite aerial athletes offer considerable insight into the constraints and demands on animal design for maximal aerobic capacity. Additionally, the similarities shown between the different phyla suggest the existence of common mechanisms and limitations in metabolic and mechanical performance. Insects in particular offer a number of advantages in pursuing questions such as the cause of the allometric scaling of muscle efficiency; this scaling can be examined within families, genera, or species with the additional benefit that insect muscles also perform well in vitro.

Preclinical abortions and endometriosis.

A single human chorionic gonadotropin determination was performed in 786 infertile women during the late luteal phase to determine the frequencies of preclinical abortions and whether the frequency was increased in women with endometriosis. Thirty-seven pregnancies (4.7% of cycles) were identified, of which six were classified as preclinical abortions (0.8%). In women with endometriosis, the frequency of preclinical abortions was 0.9% and was not statistically different from other infertile subgroups. This study suggests that preclinical abortions are not cause of infertility in either an infertile population as a whole or in the subgroup of women with endometriosis.

Power and efficiency of insect flight muscle.

The efficiency and mechanical power output of insect flight muscle have been estimated from a study of hovering flight. The maximum power output, calculated from the muscle properties, is adequate for the aerodynamic power requirements. However, the power output is insufficient to oscillate the wing mass as well unless there is good elastic storage of the inertial energy, and this is consistent with reports of elastic components in the flight system. A comparison of the mechanical power output with the metabolic power input to the flight muscles suggests that the muscle efficiency is quite low: less than 10%.