Increased Wingbeat Frequency Research Articles

Domestic egg-laying hens, Gallus gallus domesticus, do not modulate flapping flight performance in response to wing condition.

Wild birds modulate wing and whole-body kinematics to adjust their flight patterns and trajectories when wing loading increases flight power requirements. Domestic chickens (Gallus gallus domesticus) in backyards and farms exhibit feather loss, naturally high wing loading, and limited flight capabilities. Yet, housing chickens in aviaries requires birds to navigate three-dimensional spaces to access resources. To understand the impact of feather loss on laying hens' flight capabilities, we symmetrically clipped the primary and secondary feathers before measuring wing and whole-body kinematics during descent from a 1.5 m platform. We expected birds to compensate for increased wing loading by increasing wingbeat frequency, amplitude and angular velocity. Otherwise, we expected to observe an increase in descent velocity and angle and an increase in vertical acceleration. Feather clipping had a significant effect on descent velocity, descent angle and horizontal acceleration. Half-clipped hens had lower descent velocity and angle than full-clipped hens, and unclipped hens had the highest horizontal acceleration. All hens landed with a velocity two to three times greater than in bird species that are adept fliers. Our results suggest that intact laying hens operate at the maximal power output supported by their anatomy and are at the limit of their ability to control flight trajectory.

Sublethal neonicotinoid exposure attenuates the effects of electromagnetic fields on honey bee flight and learning

Many environmental stressors are currently implicated in the decline of flying insects in general, and important pollinator species such as the honey bee. Recent studies have shown that extremely low frequency electromagnetic fields (ELF EMFs) affect many aspects of insect behaviour and cognition which raises the potential that ELF EMFs could interact with other environmental stressors such as neonicotinoid insecticides to have even greater impact on the decline of flying insects. Here we analysed the effects of individual and combined exposure of the honey bee to 50 Hz EMFs and sublethal exposure to clothianidin, in a tethered flight assay and an olfactory appetitive learning assay.Clothianidin was significantly toxic to bees and exposure to field-realistic levels (2.00 ng/bee) led to 25% mortality. Exposure to ELF EMFs alone led to a significant increase in wingbeat frequency at levels above 100 µT. Prior exposure to clothianidin attenuated the effects of EMFs on wingbeat frequency. Exposure to EMFs alone reduced learning of a proboscis extension reflex (PER). Prior exposure to low doses of clothianidin attenuated the effects of EMFs on PER.These results indicate no evidence of synergy between clothianidin and ELF EMFs as environmental stressors but suggest the potential for EMFs to affect the same susceptible fraction of the bee population that have been affected by clothianidin. Results lay the foundation to further explore the interactions of ELF EMFs with other environmental stressors and consider the key factors that may make bees susceptible to ELF EMFs.

Wing damage affects flight kinematics but not flower tracking performance in hummingbird hawkmoths.

Wing integrity is crucial to the many insect species that spend distinct portions of their life in flight. How insects cope with the consequences of wing damage is therefore a central question when studying how robust flight performance is possible with such fragile chitinous wings. It has been shown in a variety of insect species that the loss in lift-force production resulting from wing damage is generally compensated by an increase in wing beat frequency rather than amplitude. The consequences of wing damage for flight performance, however, are less well understood, and vary considerably between species and behavioural tasks. One hypothesis reconciling the varying results is that wing damage might affect fast flight manoeuvres with high acceleration, but not slower ones. To test this hypothesis, we investigated the effect of wing damage on the manoeuvrability of hummingbird hawkmoths (Macroglossum stellatarum) tracking a motorised flower. This assay allowed us to sample a range of movements at different temporal frequencies, and thus assess whether wing damage affected faster or slower flight manoeuvres. We show that hummingbird hawkmoths compensate for the loss in lift force mainly by increasing wing beat amplitude, yet with a significant contribution of wing beat frequency. We did not observe any effects of wing damage on flight manoeuvrability at either high or low temporal frequencies.

Fungicide suppression of flight performance in the honeybee (Apis mellifera) and its amelioration by quercetin.

As a managed agricultural pollinator, the western honeybee Apis mellifera frequently encounters agrochemicals as contaminants of nectar and pollen. One such contaminant, the fungicide boscalid, is applied at bloom in orchards for fungal floral pathogen control. As an inhibitor of complex II in the mitochondrial electron transport chain of fungi, boscalid can potentially interfere with high energy-demanding activities of bees, including flight. We designed an indoor flight treadmill to evaluate impacts of ingesting boscalid and/or quercetin, a ubiquitous phytochemical in bee food that also affects mitochondrial respiration. Boscalid reduced the wingbeat frequencies of foragers during flight but did not alter the duration of flight. At the colony level, boscalid ingestion may thereby affect overall health by reducing forager efficiency. The consumption of quercetin, by contrast, led to higher adenosine triphosphate levels in flight muscles and a higher wingbeat frequency. Consuming the two compounds together increased wingbeat frequency, demonstrating a hitherto unrecognized mechanism by which dietary phytochemicals may act to ameliorate toxic effects of pesticides to promote honeybee health. In carrying out this work, we also introduce two methodological improvements for use in testing for pesticide effects on flight capacity-a 'force-feeding' to standardize flight fuel supply and a novel indoor flight treadmill.

Birds invest wingbeats to keep a steady head and reap the ultimate benefits of flying together.

Flapping flight is the most energetically demanding form of sustained forwards locomotion that vertebrates perform. Flock dynamics therefore have significant implications for energy expenditure. Despite this, no studies have quantified the biomechanical consequences of flying in a cluster flock or pair relative to flying solo. Here, we compared the flight characteristics of homing pigeons (Columba livia) flying solo and in pairs released from a site 7 km from home, using high-precision 5 Hz global positioning system (GPS) and 200 Hz tri-axial accelerometer bio-loggers. As expected, paired individuals benefitted from improved homing route accuracy, which reduced flight distance by 7% and time by 9%. However, realising these navigational gains involved substantial changes in flight kinematics and energetics. Both individuals in a pair increased their wingbeat frequency by 18% by decreasing the duration of their upstroke. This sharp increase in wingbeat frequency caused just a 3% increase in airspeed but reduced the oscillatory displacement of the body by 22%, which we hypothesise relates to an increased requirement for visual stability and manoeuvrability when flying in a flock or pair. The combination of the increase in airspeed and a higher wingbeat frequency would result in a minimum 2.2% increase in the total aerodynamic power requirements if the wingbeats were fully optimised. Overall, the enhanced navigational performance will offset any additional energetic costs as long as the metabolic power requirements are not increased above 9%. Our results demonstrate that the increases in wingbeat frequency when flying together have previously been underestimated by an order of magnitude and force reinterpretation of their mechanistic origin. We show that, for pigeons flying in pairs, two heads are better than one but keeping a steady head necessitates energetically costly kinematics.

Pre-copula acoustic behaviour of males in the malarial mosquitoes Anopheles coluzzii and Anopheles gambiae s.s. does not contribute to reproductive isolation.

We reveal that males of two members of the Anopheles gambiae s.l. species complex, Anopheles coluzzii and Anopheles gambiae s.s. (hereafter A. gambiae), which are both malaria vectors, perform a stereotypical acoustic behaviour in response to pure tones at frequencies that encompass the frequency range of the female's flight-tones. This behaviour resembles that described for Culex quinquefasciatus and consists of phonotactic flight initiated by a steep increase in wing-beat frequency (WBF) followed by rapid frequency modulation (RFM) of WBF when in close proximity to the sound source. RFM was elicited without acoustic feedback or the presence of a live female, but it appears to be a stereotypic behaviour in the immediate lead up to copula formation. RFM is an independent and different behavioural process from harmonic convergence interactions used by male-female pairs for mate recognition at earlier stages of mating. Acoustic threshold for RFM was used to plot behavioural audiograms from free-flying A coluzzii and A gambiae males. These audiograms were almost identical (minima ∼400 Hz) and encompassed the WBF ranges of A coluzzii (378-601 Hz) and A gambiae (373-590 Hz) females, indicating that males of the two species share similar frequency tuning and range. Furthermore, no differences were found between the two species in their WBFs, RFM behaviour or harmonic convergence ratios. These results indicate that assortative mating between A coluzzii and A gambiae is unlikely to be based on male-specific acoustic behaviours during RFM. The significance of these findings in relation to possible mechanisms for assortative mating is discussed.

Wingbeat kinematics and energetics during weightlifting in hovering hummingbirds across an elevational gradient.

Hummingbirds differentially modify flight kinematics in response to the type of challenge imposed. Weightlifting is associated with increases in stroke amplitude (the angle swept by the wings) to increase the angular velocity of the wings and generate the requisite lift, but only up to 160°. Conversely, flight in hypodense air is accomplished by increasing the angular velocity of the wing through increases in wingbeat frequency and stroke amplitudes, with larger increases in amplitude than seen in weightlifting flight. The kinematic differences between these two challenges may be facilitated by the lower energetic costs associated with overcoming drag and inertial forces over the wing during hypodense flight. Thus, we hypothesized that energetic expenditure is what limits the kinematics of weightlifting flight, with lower air densities permitting increases in angular velocity at comparatively lower costs. To explore the kinematic and energetic effects of air density and weightlifting on hovering flight performance, video and respirometric recordings of weightlifting were performed on four species of hummingbirds across an elevational gradient. Contrary to our hypothesis, wingbeat frequency did not vary due to elevation. Instead, wingbeat frequency seems to increase depending on the power requirements for sustaining hovering flight. Furthermore, metabolic rates during hovering increased with angular velocity alone, independent of elevation. Thus, it appears that the differential responses to flight challenges are not driven by variation in the flight media.

A role for acoustic distortion in novel rapid frequency modulation behaviour in free-flying male mosquitoes

We describe a new stereotypical acoustic behaviour by male mosquitoes in response to the fundamental frequency of female flight tones during mating sequences. This male-specific free-flight behaviour consists of phonotactic flight beginning with a steep increase in wing-beat frequency (WBF) followed by rapid frequency modulation (RFM) of WBF in the lead up to copula formation. Male RFM behaviour involves remarkably fast changes in WBF and can be elicited without acoustic feedback or physical presence of the female. RFM features are highly consistent, even in response to artificial tones that do not carry the multi-harmonic components of natural female flight tones. Comparison between audiograms of the robust RFM behaviour and the electrical responses of the auditory Johnston's organ (JO) reveals that the male JO is tuned not to the female WBF per se but, remarkably, to the difference between the male and female WBFs. This difference is generated in the JO responses as a result of intermodulation distortion products (DPs) caused by non-linear interaction between male-female flight tones in the vibrations of the antenna. We propose that male mosquitoes rely on their own flight tones in making use of DPs to acoustically detect, locate and orientate towards flying females. We argue that the previously documented flight-tone harmonic convergence of flying male and female mosquitoes could be a consequence of WBF adjustments so that DPs generated through flight-tone interaction fall within the optimal frequency ranges for JO detection.

Shape of wing wear fails to affect load lifting in common eastern bumble bees (Bombus impatiens) with experimental wing wear

Wing wear reflects the accumulation of irreversible damage to an insect’s wings over its lifetime and this damage should influence flight performance. In the case of bumble bees, flight seems robust to variation in wing-area asymmetry and air pressure, but not to loss of wing area. However, how the pattern of wing wear affects flight performance remains unstudied. In nature, wing wear typically occurs in a ragged and haphazard pattern along the wing’s trailing margin, a shape strikingly different from the straight cut applied in past studies. In this study, we test if shape of wing wear (implemented as four distinct treatments plus a control) affects maximum load-lifting capabilities and wingbeat frequency of worker common eastern bumble bees (Bombus impatiens Cresson, 1863). We found that shape of wing wear of 171 mg bees had no detectable effect on maximum load-lifting capability (detectable effect size = 18 mg) or on wingbeat frequency (detectable effect size = 15 Hz), but that loss of wing area reduced load-lifting capability and increased wingbeat frequency. The importance of wing area in explaining the load-lifting ability of bumble bees is reinforced in this study. But, paradoxically, shape of wing wear did not detectably affect lift generation, which is determined by unsteady aerodynamic forces in these lift-reliant insects.

Hovering flight in the honeybee Apis mellifera: kinematic mechanisms for varying aerodynamic forces.

During hovering flight, animals can increase the wing velocity and therefore the net aerodynamic force per stroke by increasing wingbeat frequency, wing stroke amplitude, or both. The magnitude and orientation of aerodynamic forces are also influenced by the geometric angle of attack, timing of wing rotation, wing contact, and pattern of deviation from the primary stroke plane. Most of the kinematic data available for flying animals are average values for wing stroke amplitude and wingbeat frequency because these features are relatively easy to measure, but it is frequently suggested that the more subtle and difficult-to-measure features of wing kinematics can explain variation in force production for different flight behaviors. Here, we test this hypothesis with multicamera high-speed recording and digitization of wing kinematics of honeybees (Apis mellifera) hovering and ascending in air and hovering in a hypodense gas (heliox: 21% O2, 79% He). Bees employed low stroke amplitudes (86.7° ± 7.9°) and high wingbeat frequencies (226.8 ± 12.8 Hz) when hovering in air. When ascending in air or hovering in heliox, bees increased stroke amplitude by 30%-45%, which yielded a much higher wing tip velocity relative to that during simple hovering in air. Across the three flight conditions, there were no statistical differences in the amplitude of wing stroke deviation, minimum and stroke-averaged geometric angle of attack, maximum wing rotation velocity, or even wingbeat frequency. We employed a quasi-steady aerodynamic model to estimate the effects of wing tip velocity and geometric angle of attack on lift and drag. Lift forces were sensitive to variation in wing tip velocity, whereas drag was sensitive to both variation in wing tip velocity and angle of attack. Bees utilized kinematic patterns that did not maximize lift production but rather maintained lift-to-drag ratio. Thus, our data indicate that, at least for honeybees, the overall time course of wing angles is generally preserved and modulation of wing tip velocity is sufficient to perform a diverse set of vertical flight behaviors.

How wing kinematics affect power requirements and aerodynamic force production in a robotic bat wing

Bats display a wide variety of behaviors that require different amounts of aerodynamic force. To control and modulate aerodynamic force, bats change wing kinematics, which, in turn, may change the power required for wing motion. There are many kinematic mechanisms that bats, and other flapping animals, can use to increase aerodynamic force, e.g. increasing wingbeat frequency or amplitude. However, we do not know if there is a difference in energetic cost between these different kinematic mechanisms. To assess the relationship between mechanical power input and aerodynamic force output across different isolated kinematic parameters, we programmed a robotic bat wing to flap over a range of kinematic parameters and measured aerodynamic force and mechanical power. We systematically varied five kinematic parameters: wingbeat frequency, wingbeat amplitude, stroke plane angle, downstroke ratio, and wing folding. Kinematic values were based on observed values from free flying Cynopterus brachyotis, the species on which the robot was based. We describe how lift, thrust, and power change with increases in each kinematic variable. We compare the power costs associated with generating additional force through the four kinematic mechanisms controlled at the shoulder, and show that all four mechanisms require approximately the same power to generate a given force. This result suggests that no single parameter offers an energetic advantage over the others. Finally, we show that retracting the wing during upstroke reduces power requirements for flapping and increases net lift production, but decreases net thrust production. These results compare well with studies performed on C. brachyotis, offering insight into natural flight kinematics.

The ACTC A295S Mutation Increases Heart and Skeletal Muscle Contractile Activity in Drosophila Models of Hypertrophic Cardiomyopathy

Hypertrophic cardiomyopathy (HCM) is an autosomal dominant disease characterized by thickened heart walls, cardiac arrhythmias, myocyte disarray and interstitial fibrosis. Over 10 mutations in alpha-cardiac actin (ACTC) are associated with HCM. The ACTC Ala295Ser substitution causes a highly penetrant disease with diverse phenotypes. To investigate the in vivo consequences of the ACTC mutation we generated Drosophila lines with muscle-specific expression of A295S actin. Our unique models limit genetic diversity and pathological complexity to help resolve molecular mechanisms of diseases. For expression in the fly heart we used UAS-Act57BWT and UAS-Act57BA295S actin transgenes in conjunction with the HandGal4 cardiac-specific driver. Confocal microscopy confirmed heart-restricted expression of UAS-Act57BWTGFP and UAS-Act57BA295SGFP and co-polymerization of transgenic and endogenous actin. High-speed video microscopy and motion analysis revealed A295S actin expression significantly reduced diastolic volumes and increased systolic intervals consistent with elevated contractile properties at rest and during contraction. For indirect flight muscle (IFM)-specific actin expression, we used the IFM targeted Act88FWTand Act88FA295S transgenes, in an IFM actin null background. Muscle mechanics on heterozygous IFM fibers revealed that the A295S actin mutation increased power, the frequency of maximum power generation (fmax), and stiffness relative to control fibers. The higher fmax and stiffness are likely causes of an observed increase in wing beat frequency (WBF) of Act88FA295Sheterozygous flies. The WBF increase had a negative impact on flight ability which decreased further as mutant: wildtype actin ratios were increased. Overall, enhanced contractile activity of actin HCM cardiac myocytes and IFM fibers is consistent with the hypercontractile properties frequently associated with HCM mutations.

Hawkmoth flight stability in turbulent vortex streets

Shedding of vortices is a common phenomenon in the atmosphere over a wide range of spatial and temporal scales. However, it is unclear how these vortices of varying scales affect the flight performance of flying animals. In order to examine these interactions, we trained seven hawkmoths (Manduca sexta) (wingspan ~9 cm) to fly and feed in a wind tunnel under steady flow (controls) and in the von Kármán vortex street of vertically oriented cylinders (two different cylinders with diameters of 10 and 5 cm) at speeds of 0.5, 1 and 2 m s(-1). Cylinders were placed at distances of 5, 25 and 100 cm upstream of the moths. Moths exhibited large amplitude yaw oscillations coupled with modest oscillations in roll and pitch, and slight increases in wingbeat frequency when flying in both the near (recirculating) and middle (vortex dominated) wake regions. Wingbeat amplitude did not vary among treatments, except at 1 m s(-1) for the large cylinder. Yaw and roll oscillations were synchronized with the vortex shedding frequencies in moths flying in the wake of the large cylinder at all speeds. In contrast, yaw and pitch were synchronized with the shedding frequency of small vortices at speeds ≤1 m s(-1). Oscillations in body orientation were also substantially smaller in the small cylinder treatment when compared with the large cylinder, regardless of temporal or non-dimensional spatial scale. Moths flying in steady conditions reached a higher air speed than those flying into cylinder wakes. In general, flight effects produced by the cylinder wakes were qualitatively similar among the recirculating and vortex-dominated wake regions; the magnitude of those effects, however, declined gradually with downstream distance.

Flying in the rain: hovering performance of Anna's hummingbirds under varied precipitation

Flight in rain represents a greater challenge for smaller animals because the relative effects of water loading and drop impact are greater at reduced scales given the increased ratios of surface area to mass. Nevertheless, it is well known that small volant taxa such as hummingbirds can continue foraging even in extreme precipitation. Here, we evaluated the effect of four rain intensities (i.e. zero, light, moderate and heavy) on the hovering performance of Anna's hummingbirds (Calypte anna) under laboratory conditions. Light-to-moderate rain had only a marginal effect on flight kinematics; wingbeat frequency of individuals in moderate rain was reduced by 7 per cent relative to control conditions. By contrast, birds hovering in heavy rain adopted more horizontal body and tail positions, and also increased wingbeat frequency substantially, while reducing stroke amplitude when compared with control conditions. The ratio between peak forces produced by single drops on a wing and on a solid surface suggests that feathers can absorb associated impact forces by up to approximately 50 per cent. Remarkably, hummingbirds hovered well even under heavy precipitation (i.e. 270 mm h(-1)) with no apparent loss of control, although mechanical power output assuming perfect and zero storage of elastic energy was estimated to be about 9 and 57 per cent higher, respectively, compared with normal hovering.

Kinematic plasticity during flight in fruit bats: individual variability in response to loading.

All bats experience daily and seasonal fluctuation in body mass. An increase in mass requires changes in flight kinematics to produce the extra lift necessary to compensate for increased weight. How bats modify their kinematics to increase lift, however, is not well understood. In this study, we investigated the effect of a 20% increase in mass on flight kinematics for Cynopterus brachyotis, the lesser dog-faced fruit bat. We reconstructed the 3D wing kinematics and how they changed with the additional mass. Bats showed a marked change in wing kinematics in response to loading, but changes varied among individuals. Each bat adjusted a different combination of kinematic parameters to increase lift, indicating that aerodynamic force generation can be modulated in multiple ways. Two main kinematic strategies were distinguished: bats either changed the motion of the wings by primarily increasing wingbeat frequency, or changed the configuration of the wings by increasing wing area and camber. The complex, individual-dependent response to increased loading in our bats points to an underappreciated aspect of locomotor control, in which the inherent complexity of the biomechanical system allows for kinematic plasticity. The kinematic plasticity and functional redundancy observed in bat flight can have evolutionary consequences, such as an increase potential for morphological and kinematic diversification due to weakened locomotor trade-offs.

Olfactory modulation of flight in Drosophila is sensitive, selective and rapid

Freely flying Drosophila melanogaster respond to odors by increasing their flight speed and turning upwind. Both these flight behaviors can be recapitulated in a tethered fly, which permits the odor stimulus to be precisely controlled. In this study, we investigated the relationship between these behaviors and odor-evoked activity in primary sensory neurons. First, we verified that these behaviors are abolished by mutations that silence olfactory receptor neurons (ORNs). We also found that antennal mechanosensors in Johnston's organ are required to guide upwind turns. Flight responses to an odor depend on the identity of the ORNs that are active, meaning that these behaviors involve odor discrimination and not just odor detection. Flight modulation can begin rapidly (within about 85 ms) after the onset of olfactory transduction. Moreover, just a handful of spikes in a single ORN type is sufficient to trigger these behaviors. Finally, we found that the upwind turn is triggered independently from the increase in wingbeat frequency, implying that ORN signals diverge to activate two independent and parallel motor commands. Together, our results show that odor-evoked flight modulations are rapid and sensitive responses to specific patterns of sensory neuron activity. This makes these behaviors a useful paradigm for studying the relationship between sensory neuron activity and behavioral decision-making in a simple and genetically tractable organism.

Neuromuscular control of wingbeat kinematics in Anna's hummingbirds (Calypte anna)

Hummingbirds can maintain the highest wingbeat frequencies of any flying vertebrate - a feat accomplished by the large pectoral muscles that power the wing strokes. An unusual feature of these muscles is that they are activated by one or a few spikes per cycle as revealed by electromyogram recordings (EMGs). The relatively simple nature of this activation pattern provides an opportunity to understand how motor units are recruited to modulate limb kinematics. Hummingbirds made to fly in low-density air responded by moderately increasing wingbeat frequency and substantially increasing the wing stroke amplitude as compared with flight in normal air. There was little change in the number of spikes per EMG burst in the pectoralis major muscle between flight in normal and low-density heliox (mean=1.4 spikes cycle(-1)). However the spike amplitude, which we take to be an indication of the number of active motor units, increased in concert with the wing stroke amplitude, 1.7 times the value in air. We also challenged the hummingbirds using transient load lifting to elicit maximum burst performance. During maximum load lifting, both wing stroke amplitude and wingbeat frequency increased substantially above those values during hovering flight. The number of spikes per EMG burst increased to a mean of 3.3 per cycle, and the maximum spike amplitude increased to approximately 1.6 times those values during flight in heliox. These results suggest that hummingbirds recruit additional motor units (spatial recruitment) to regulate wing stroke amplitude but that temporal recruitment is also required to maintain maximum stroke amplitude at the highest wingbeat frequencies.

Hovering Energetics and Thermal Balance in Anna’s Hummingbirds (Calypte anna)

We studied the energetics of hover-feeding Anna's hummingbirds, using three different simultaneous techniques: heat loss as estimated via thermal imaging, metabolic rate as measured at a feeder mask using flow-through respirometry, and aerodynamic power estimated from wingbeat kinematic data. These three methods yielded comparable estimates of power output at ambient air temperatures ranging from 18 degrees to 26 degrees C, whereas heat imbalance at higher air temperatures (up to 34 degrees C) suggested loss by mechanisms other than convection and radiation from the body, such as evaporative cooling and enthalpy rise associated with exhaled air and excreted water and convective heat loss from the patagia. Hummingbirds increased wingbeat frequency and decreased stroke amplitude as air temperature increased, but overall muscle efficiency was found to be approximately constant over the experimental range of air temperatures.

The scaling of myofibrillar actomyosin ATPase activity in apid bee flight muscle in relation to hovering flight energetics

For all types of locomotion, the overall efficiency with which chemical energy is converted into mechanical work increases with increasing body size. In order to gain insight into the determinants of the scaling of overall efficiency, we measured the scaling of the rate of ATP utilisation during cyclical contractions using glycerinated fibres from the dorsolongitudinal flight muscle of several species of apid bees, covering a ninefold range in body mass. The efficiency of ATP utilisation by the crossbridges is one of the stages that determines the overall efficiency of locomotion. The mechanochemical coefficient was calculated from the ratio of the net power output to the rate of ATP hydrolysis and ranged from 6.5 to 9.7 kJ mol(-1) ATP. The corresponding gross myofibrillar efficiency was 15-23%, increasing concomitantly with body mass (M(b)) and decreasing with increasing wingbeat frequency (n) and scaling as M(b)(0.184) and n(-1.168) in bumblebees and as M(b)(0.153) and n(-0.482) in euglossine bees. Overall efficiency of hovering in bumblebees and euglossine bees was calculated using previously published metabolic power data and revised estimates of the mechanical power output to take into account the drag due to the leading edge vortex that has not been included in previous models. The scaling of overall efficiency of hovering flight in apid bees was not as pronounced as the scaling of myofibrillar efficiency. Therefore the scaling of myofibrillar efficiency with body mass (or frequency) only explained part of the scaling of overall efficiency, and it is likely that the efficiency of other steps in the transduction of chemical energy into mechanical work (e.g. the efficiency of mitochondrial oxidative recovery) may also scale with body mass.

Wing beat kinematics of a nectar-feeding bat,Glossophaga soricina,flying at different flight speeds and Strouhal numbers

High-speed film analysis showed that the wing beat kinematics in Glossophaga soricina change gradually with increasing flight speed, indicating that there is no sudden gait change at any particular, critical, flight speed. The flight of two adult specimens was studied over a range of flight speeds (1.23-7.52 ms(-1)) in a 30 m long flight tunnel. During the upstroke in hovering and slow flight there is a tip-reversal or supination of the handwings, which thus produce a backward flick. This backward motion successively disappears at speeds V approximately 3.2 ms(-1), above which the wingtip path becomes more vertical or directed upwards-forwards relative to the still air (the stroke plane angle increasing with flight speed as alpha=44.8V(0.29)). We found no correlations between either span ratio SR (the ratio of the wing span on the upstroke to that on the downstroke) and V, or downstroke ratio (the duration of the downstroke divided by the total stroke period) and V. On the other hand, SR decreases significantly with increasing wing beat frequency f, SR proportional to f(-0.40). The Strouhal number (St=f x amplitude/V), a dimensionless parameter describing oscillating flow mechanisms and being a predictor of the unsteadiness of the flow, decreases with the speed as St proportional to V(-1.37). Close to the theoretical minimum power speed (4-6 m s(-1)) G. soricina operates with a Strouhal number in the region 0.17<St<0.22, which is associated with efficient lift and thrust production. At slower speeds, 3.4-4 m s(-1), St is 0.25-0.4, which is still within the favourable region. But at speeds below 3 m s(-1) St becomes higher (0.5<St<0.68), indicating that unsteady effects become important, with unfavourable lift and thrust production as a result. Only at these speeds do the bats perform the backward flick during the upstroke, which may produce thrust. This may serve as a compensation in some bats and birds to increase aerodynamic performance.