Locust flight initiation: a comparison of normal and artificial release

Flight initiation was analysed in the migratory locust, Locusta migratoria L. using conventional and high‐speed video systems. Flight was initiated by three methods: normal jump, free fall and controlled catapult. Parameters evaluated were time from release to wing‐opening, body angle, speed, height gain, wing‐opening, initial wing‐beat frequency. Fastest wing‐opening occurred following a normal jump. A catapult device allowed manipulation of the launching parameters: speed and angles at launching. It appeared that in an artificially catapulted start there was a minimum speed (v > 0.75 m/s) required to initiate flight. However, under free‐fall conditions a mean speed of v = 0.6 m/s at wing‐opening was observed. When the different parameters of the controlled catapult start were equal to those for normal jump then the time to wing‐opening was found to be extended for the catapult launch. However, other parameters were not affected, occasionally even a ‘kick in air’ was observed. The catapult launches indicated that within about three wing‐beat cycles the animals showed active flight, with positive lift and constant or increasing speed, compared to a ballistic trajectory. Our results indicate that a controlled catapult device will prove useful to the study of sensory and central processes underlying free flight initiation.     

Flight costs of long,sexually selected tails in hummingbirds

The elongated tails adorning many male birds have traditionally been thought to degrade flight performance by increasing body drag. However, aerodynamic interactions between the body and tail can be substantial in some contexts, and a short tail may actually reduce rather than increase overall drag. To test how tail length affects flight performance, we manipulated the tails of Anna''s hummingbirds (Calypte anna) by increasing their length with the greatly elongated tail streamers of the red-billed streamertail (Trochilus polytmus) and reducing their length by removing first the rectrices and then the entire tail (i.e. all rectrices and tail covert feathers). Flight performance was measured in a wind tunnel by measuring (i) the maximum forward speed at which the birds could fly and (ii) the metabolic cost of flight while flying at airspeeds from 0 to 14 m s⁻¹. We found a significant interaction effect between tail treatment and airspeed: an elongated tail increased the metabolic cost of flight by up to 11 per cent, and this effect was strongest at higher flight speeds. Maximum flight speed was concomitantly reduced by 3.4 per cent. Also, removing the entire tail decreased maximum flight speed by 2 per cent, suggesting beneficial aerodynamic effects for tails of normal length. The effects of elongation are thus subtle and airspeed-specific, suggesting that diversity in avian tail morphology is associated with only modest flight costs.     

Development of flight behaviour in maturing adults of Locusta migratoria: I. Flight performance and wing-stroke parameters

In tethered Locusta migratoria suspended from a flight balance, flight performance, wing-stroke frequency, stroke angle, and stroke plane angle were studied throughout adult life. No correlation between flight performance and age was found in adults older than 2 days. During continuous flight in locusts of all ages the wing-stroke frequency and the wing-stroke angle of both wings decreases, and the wing-stroke plane angle (forewing) increases slightly. Within 2 weeks of adult life the wing-stroke frequency increases by a factor of ca. 2, whereas the wing-stroke angles and the stroke plane angles remain constant.     

Kinematic compensation for wing loss in flying damselflies

Flying insects can tolerate substantial wing wear before their ability to fly is entirely compromised. In order to keep flying with damaged wings, the entire flight apparatus needs to adjust its action to compensate for the reduced aerodynamic force and to balance the asymmetries in area and shape of the damaged wings. While several studies have shown that damaged wings change their flapping kinematics in response to partial loss of wing area, it is unclear how, in insects with four separate wings, the remaining three wings compensate for the loss of a fourth wing. We used high-speed video of flying blue-tailed damselflies (Ischnura elegans) to identify the wingbeat kinematics of the two wing pairs and compared it to the flapping kinematics after one of the hindwings was artificially removed. The insects remained capable of flying and precise maneuvering using only three wings. To compensate for the reduction in lift, they increased flapping frequency by 18 ± 15.4% on average. To achieve steady straight flight, the remaining intact hindwing reduced its flapping amplitude while the forewings changed their stroke plane angle so that the forewing of the manipulated side flapped at a shallower stroke plane angle. In addition, the angular position of the stroke reversal points became asymmetrical. When the wingbeat amplitude and frequency of the three wings were used as input in a simple aerodynamic model, the estimation of total aerodynamic force was not significantly different (paired t-test, p = 0.73) from the force produced by the four wings during normal flight. Thus, the removal of one wing resulted in adjustments of the motions of the remaining three wings, exemplifying the precision and plasticity of coordination between the operational wings. Such coordination is vital for precise maneuvering during normal flight but it also provides the means to maintain flight when some of the wings are severely damaged.     

Flight behavior of the bean bug,Riptortus clavatus (Thunberg) (Hemiptera: Alydidae), relative to adult age,sex and season

Studies related to the flight behavior of the bean bug, Riptortus clavatus, an insect pest of soybean, provide information, which can aid the development of management tactics. R. clavatus flight activities were determined using the tethered flight technique in the laboratory and a field study. We compared the flight parameters (flight distance, duration, speed and frequency) of laboratory-reared mated or unmated females and males of different adult age groups, and field-collected females and males from different seasons in a year. Mating effect was only significant for flight frequency, which was higher for unmated adults. Only the flight frequency was different between females and males. Among age groups, 25 d old individuals were shown to have higher flight parameters while the 35 or 45 d old groups showed the lower flight. Among the season, flight activities decreased as the season progressed. The flight distance of R. clavatus in a day was estimated to be 1.6–5.1 km with an average speed of 0.8 m/s from the laboratory data. However, from the field study, it was shown that bean bugs flew a 45–54 m distance with a flight speed of 3.0–3.6 m/s for the first single flight. The second flight was much shorter and slower. We discuss the possible difference of flight parameters between the laboratory and field studies with features of flight mill, variable field conditions and host plant finding behaviors. Further study on flight behavior may provide a better understanding of R. clavatus biology which may guide proper management.     

Flow visualization and unsteady aerodynamics in the flight of the hawkmoth,Manduca sexta

The aerodynamic mechanisms employed durng the flight of the hawkmoth, Manduca sexta, have been investigated through smoke visualization studies with tethered moths. Details of the flow around the wings and of the overall wake structure were recorded as stereophotographs and high-speed video sequences. The changes in flow which accompanied increases in flight speed from 0.4 to 5.7 m s^-1 were analysed. The wake consists of an alternating series of horizontal and vertical vortex rings which are generated by successive down- and upstrokes, respectively. The downstroke produces significantly more lift than the upstroke due to a leading-edge vortex which is stabilized by a radia flow moving out towards the wingtip. The leading-edge vortex grew in size with increasing forward flight velocity. Such a phenomenon is proposed as a likely mechanism for lift enhancement in many insect groups. During supination, vorticity is shed from the leading edge as postulated in the ''flex'' mechanism. This vorticity would enhance upstroke lift if it was recaptured diring subsequent translation, but it is not. Instead, the vorticity is left behind and the upstroke circulation builds up slowly. A small jet provides additional thrust as the trailing edges approach at the end of the upstroke. The stereophotographs also suggest that the bound circulation may not be reversed between half strokes at the fastest flight speeds.     

Flight responses by a migratory soaring raptor to changing meteorological conditions

Soaring birds that undertake long-distance migration should develop strategies to minimize the energetic costs of endurance flight. This is relevant because condition upon completion of migration has direct consequences for fecundity, fitness and thus, demography. Therefore, strong evolutionary pressures are expected for energy minimization tactics linked to weather and topography. Importantly, the minute-by-minute mechanisms birds use to subsidize migration in variable weather are largely unknown, in large part because of the technological limitations in studying detailed long-distance bird flight. Here, we show golden eagle (Aquila chrysaetos) migratory response to changing meteorological conditions as monitored by high-resolution telemetry. In contrast to expectations, responses to meteorological variability were stereotyped across the 10 individuals studied. Eagles reacted to increased wind speed by using more orographic lift and less thermal lift. Concomitantly, as use of thermals decreased, variation in flight speed and altitude also decreased. These results demonstrate how soaring migrant birds can minimize energetic expenditures, they show the context for avian decisions and choices of specific instantaneous flight mechanisms and they have important implications for design of bird-friendly wind energy.     

Flight capability and flight motor pattern in a sedentary South African grasshopper, Phymateus morbillosus– a comparison with migratory species

Abstract Among the Acridoidea, not all species are strong fliers. We have examined the possible causes for loss of flight in a species with a reduced flight system, the South African grasshopper, Phymateus morbillosus (L.). This is a sedentary species that, in the field, displays only marginal flight (in males) or no flight (in females). In a wind tunnel, however, this species can be stimulated to perform flight movements for a short time. In the present study, several morphometric parameters and aspects of the flight motor output have been examined. The data are compared with those of the migratory locust species ( Locusta migratoria L. and Schistocerca gregaria Forsk.). Phymateus morbillosus can exhibit the typical flight motor pattern for short periods of up to 1 min. Morphometric data and wing-beat frequency in P. morbillosus are similar to that of other members of this insect group. However, female specimens of P. morbillosus are too heavy to lift themselves for active flight. We assume that females of this species invest in the augmentation of reproduction rather than investing in the flight system.     

Flight speeds of swifts (Apus apus): seasonal differences smaller than expected

We have studied the nocturnal flight behaviour of the common swift (Apus apus L.), by the use of a tracking radar. Birds were tracked from Lund University in southern Sweden during spring migration, summer roosting flights and autumn migration. Flight speeds were compared with predictions from flight mechanical and optimal migration theories. During spring, flight speeds were predicted to be higher than during both summer and autumn due to time restriction. In such cases, birds fly at a flight speed that maximizes the overall speed of migration. For summer roosting flights, speeds were predicted to be lower than during both spring and autumn since the predicted flight speed is the minimum power speed that involves the lowest energy consumption per unit time. During autumn, we expected flight speeds to be higher than during summer but lower than during spring since the expected flight speed is the maximum range speed, which involves the lowest energy consumption per unit distance. Flight speeds during spring were indeed higher than during both summer and autumn, which indicates time-selected spring migration. Speeds during autumn migration were very similar to those recorded during summer roosting flights. The general result shows that swifts change their flight speed between different flight behaviours to a smaller extent than expected. Furthermore, the difference between flight speeds during migration and roosting among swifts was found to be less pronounced than previously recorded.     

Morphological and kinematic basis of the hummingbird flight stroke: scaling of flight muscle transmission ratio

Hummingbirds (Trochilidae) are widely known for their insect-like flight strokes characterized by high wing beat frequency, small muscle strains and a highly supinated wing orientation during upstroke that allows for lift production in both halves of the stroke cycle. Here, we show that hummingbirds achieve these functional traits within the limits imposed by a vertebrate endoskeleton and muscle physiology by accentuating a wing inversion mechanism found in other birds and using long-axis rotational movement of the humerus. In hummingbirds, long-axis rotation of the humerus creates additional wing translational movement, supplementing that produced by the humeral elevation and depression movements of a typical avian flight stroke. This adaptation increases the wing-to-muscle-transmission ratio, and is emblematic of a widespread scaling trend among flying animals whereby wing-to-muscle-transmission ratio varies inversely with mass, allowing animals of vastly different sizes to accommodate aerodynamic, biomechanical and physiological constraints on muscle-powered flapping flight.     

New modeling approach for bounding flight in birds

A new modeling approach is presented which accounts for the unsteady motion features and dynamics characteristics of bounding flight. For this purpose, a realistic mathematical model is developed to describe the flight dynamics of a bird with regard to a motion which comprises flapping and bound phases involving acceleration and deceleration as well as, simultaneously, pull-up and push-down maneuvers. Furthermore, a mathematical optimization method is used for determining that bounding flight mode which yields the minimum energy expenditure per range. Thus, it can be shown to what extent bounding flight is aerodynamically superior to continuous flapping flight, yielding a reduction in the energy expenditure in the speed range practically above the maximum range speed. Moreover, the role of the body lift for the efficiency of bounding flight is identified and quantified. Introducing an appropriate non-dimensionalization of the relations describing the bird’s flight dynamics, results of generally valid nature are derived for the addressed items.     

Effects of flight behaviour on body temperature and kinematics during inter-male mate competition in the solitary desert bee Centris pallida

Abstract. Body temperatures and kinematics are measured for male Centris pallida bees engaged in a variety of flight behaviours (hovering, patrolling, pursuit) at a nest aggregation site in the Sonoran Desert. The aim of the study is to test for evidence of thermoregulatory variation in convective heat loss and metabolic heat production and to assess the mechanisms of acceleration and forward flight in field conditions. Patrolling males have slightly (1–3 °C) cooler body temperatures than hoverers, despite similar wingbeat frequencies and larger body masses, suggesting that convective heat loss is likely to be greater during patrolling flight than during hovering. Comparisons of thorax and head temperature as a function of air temperature (T_a) indicate that C. pallida males are thermoregulating the head by increasing heat transfer from the thorax to the head at cool T_a. During patrolling flight and hovering, wingbeat frequency significantly decreases as T_a increases, indicating that variation in metabolic heat production contributes to thermal stability during these behaviours, as has been previously demonstrated for this species during flight in a metabolic chamber. However, wingbeat frequency during brief (1–2 s) pursuits is significantly higher than during other flight behaviours and independent of T_a. Unlike most other hovering insects, C. pallida males hover with extremely inclined stroke plane angles and nearly horizontal body angles, suggesting that its ability to vary flight speed depends on changes in wingbeat frequency and other kinematic mechanisms that are not yet described.     

Wing‐beat characteristics of birds recorded with tracking radar and cine camera

This study presents wing‐beat frequency data measured mainly by radar, complemented by video and cinematic recordings, for 153 western Palaearctic and two African species. Data on a further 45 Palaearctic species from other sources are provided in an electronic appendix. For 41 species with passerine‐type flight, the duration of flapping and pausing phases is given. The graphical presentations of frequency ranges and wing‐beat patterns show within‐species variation and allow easy comparison between species, taxonomic groups and types of flight. Wing‐beat frequency is described by Pennycuick (J. Exp. Biol. 2001; 204: 3283–3294) as a function of body‐mass, wing‐span, wing‐area, gravity and air density; for birds with passerine‐type flight the power‐fraction has also to be considered. We tested Pennycuick’s general allometric model and estimated the coefficients based on our data. The general model explained a high proportion of variation in wing‐beat frequency and the coefficients differed only slightly from Pennycuick’s original values. Modelling continuous‐flapping flyers alone resulted in coefficients not different from those predicted (within 95% intervals). Doing so for passerine‐type birds resulted in a model with non‐significant contributions of body‐mass and wing‐span to the model. This was mainly due to the very high correlation between body‐mass, wing‐span and wing‐area, revealing similar relative scaling properties within this flight type. However, wing‐beat frequency increased less than expected with respect to power‐fraction, indicating that the drop in flight level during the non‐flapping phases, compensated by the factor (g/q)^0.5 in Pennycuick’s model, is smaller than presumed. This may be due to lift produced by the body during the bounding phase or by only partial folding of the wings.     

Substrate utilization and flight speed during tethered flight in the locust

Mature laboratory locusts normally exhibit a characteristic pattern of change in flight speed with time. They fly at high speed for the first few minutes, during which carbohydrate forms the major fuel, but then slow to a cruising speed when lipid is used almost exclusively. Locusts flown for 30 min, rested for 2hr, and then reflown, exhibit an identical pattern of flight, even though they oxidise only half the amount of carbohydrate used in the first flight. The injection of adipokinetic hormone before the first flight elicits a low initial flight speed for 10 to 15 min but then the locusts accelerate to a constant higher speed. The injection of hormone before the second flight, when blood lipid levels are already high, reduces the utilization of carbohydrate by the flight muscles dramatically but results in constant high-speed flight.     

Morphology, Velocity, and Intermittent Flight in Birds

Body size, pectoralis composition, aspect ratio of the wing,and forward speed affect the use of intermittent flight in birds.During intermittent non-flapping phases, birds extend theirwings and glide or flex their wings and bound. The pectoralismuscle is active during glides but not during bounds; activityin other primary flight muscles is variable. Mechanical power,altitude, and velocity vary among wingbeats in flapping phases;associated with this variation are changes in neuromuscularrecruitment, wingbeat frequency, amplitude, and gait. Speciesof intermediate body mass (35–158 g) tend to flap-glideat slower speeds and flap-bound at faster speeds, regardlessof the aspect ratio of their wings. Such behavior may reducemechanical power output relative to continuous flapping. Smallerspecies (<20 g) with wings of low aspect ratio may flap-boundat all speeds, yet existing models do not predict an aerodynamicadvantage for the flight style at slow speeds. The behaviorof these species appears to be due to wing shape rather thanpectoralis physiology. As body size increases among species,percent time spent flapping increases, and birds much largerthan 300 g do not flap-bound. This pattern may be explainedby adverse scaling of mass-specific power or lift per unit poweroutput available from flight muscles. The size limit for theability to bound intermittently may be offset somewhat by thescaling of pectoralis composition. The percentage of time spentflapping during intermittent flight also varies according toflight speed.     

Metabolic costs of avian flight in relation to flight velocity: a study in Rose Coloured Starlings (Sturnus roseus, Linnaeus)

The metabolic costs of flight at a natural range of speeds were investigated in Rose Coloured Starlings (Sturnus roseus, Linnaeus) using doubly labelled water. Eight birds flew repeatedly and unrestrained for bouts of 6 h at speeds from 9 to 14 m s⁻¹ in a low-turbulence wind tunnel, corresponding to travel distances between 200 and 300 km, respectively. This represents the widest speed range where we could obtain voluntarily sustained flights. From a subset of these flights, data on the wing beat frequency (WBF) and intermittent flight behaviour were obtained. Over the range of speeds that were tested, flight costs did not change with velocity and were on an average 8.17±0.64 W or 114 W kg⁻¹. Body mass was the only parameter with a significant (positive) effect on flight costs, which can be described as EE_f=0.741 M ^0.554. WBF changed slightly with speed, but correlated better with body mass. Birds showed both types of intermittent flight, undulating and bounding, but their frequencies did not systematically change with flight speed.     

Dynamics of energy substrates in the haemolymph of Locusta migratoria during flight

In the two-fuel system for flight of the migratory locust, the haemolymph carbohydrate concentration falls during flight periods of up to 1 hr, the decrease being greater in case the pre-flight carbohydrate level is higher. The increase in the lipid concentration from the onset of flight is virtually independent of the initial lipid concentration. Flight intensity affects these changes in substrate concentrations: the carbohydrate level decreases more rapidly if flight speed is higher, whereas the increase in lipid concentration is delayed at higher flight speeds. Respiratory carbon dioxide production is elevated rapidly during flight and reaches over eight times the resting level. From the rate of ¹⁴CO₂ production after labelling of the haemolymph diglyceride pool it is concluded that diglycerides contribute to providing the energy for flight from the earliest stage of flying activity; diglyceride oxidation increases until maximum utilization is attained after some 45 min of flight. The decline in haemolymph carbohydrate concentration due to flying activity results in a decrease of haemolymph osmolarity. Free amino acids, particularly taurine, increase markedly in the haemolymph during flight; yet their concentration only partially counterbalances the fall in haemolymph osmolarity.     

Stabilization of altitude and speed in tethered flying gypsy moth males: influence of (+) and (-)-disparlure

ABSTRACT. Changes in lift and thrust were elicited in tethered male gypsy moths, Lymantria dispar L. (Lepidoptera, Lymantriidae), by visual pattern elements moving radially either towards or from the point directly beneath their body, if the sex-pheromone, (+)-disparlure, was present. The sign of these changes was such as to counteract the pattern movements, which were generated by a rotating spiral beneath the moth. By restricting the area of spiral visible to the moth to either transverse or longitudinal sectors, flight altitude was affected by the centrifugal/centripetal movements in the lateral sectors, whereas flight speed was affected by those in the frontal sector. It is deduced that in free flight these compensatory reactions are responsible for the stabilization of flight altitude and speed, respectively. Surprisingly, without pheromone present these responses were usually not detectable: a wide range of flight altitude and speed was then observed. In the presence of (+)-disparlure, however, these responses were always strongly pronounced, the animal keeping within a narrow range of speed and altitude. These compensatory reactions were blocked by the attraction-inhibiting (-)-disparlure if presented in racemic mixture with the (+) form: the range of speed and altitude shown by the moth was then the same as without any pheromone. Under closed-loop conditions, the mean flight speed was reduced by the racemic mixture as well as by (+)-disparlure alone, however.     

Dynamic Modeling, Testing, and Stability Analysis of an Ornithoptic Blimp

In order to study omithopter flight and to improve a dynamic model of flapping propulsion,a series of tests are conducted on a flapping-wing blimp.The blimp is designed and constructed from mylar plastic and balsa wood as a test platform for aerodynamics and flight dynamics.The blimp,2.3 meters long and 420 gram mass,is propelled by its flapping wings.Due to buoyancy the wings have no lift requirement so that the distinction between lift and propulsion can be analyzed in a flight platform at low flight speeds.The blimp is tested using a Vicon motion tracking system and various initial conditions are tested including accelerating flight from standstill,decelerating from an initial speed higher than its steady state,and from its steady-state speed but disturbed in pitch angle.Test results are used to estimate parameters in a coupled quasi-steady aerodynamics/Newtonian flight dynamics model.This model is then analyzed using Floquet theory to determine local dynamic modes and stability.It is concluded that the dynamic model adequately describes the vehicle's nonlinear behavior near the steady-state velocity and that the vehicle's linearized modes are akin to those of a fixed-wing aircraft.     

METHOPRENE INFLUENCES REPRODUCTION AND FLIGHT CAPACITY IN ADULTS OF THE RICE LEAF ROLLER,Cnaphalocrocis Medinalis (GUENỂE) (LEPIDOPTERA: PYRALIDAE)

Juvenile hormone (JH) influences many aspects of insect biology, including oogenesis‐flight syndrome tradeoffs between migration and reproduction. Drawing on studies of many migratory insects, we posed the hypothesis that JH influences migratory capacity and oogenesis in the rice leaf roller, Cnaphalocrocis medinalis. We treated adults moths (days 1, 2 and 3 postemergence) with the JH analog (JHA), methoprene, and then recorded the influences of JHA treatments on reproduction. JHA treatment on day 1 postemergence, but not on the other days, shortened the preoviposition period, although JHA did not influence total fecundity, oviposition period, or longevity. We infer day 1 postemergence is the JH‐sensitive stage to influence reproduction. Therefore, we treated moths on day 1 postemergence with JHA and recorded flight capacity, flight muscle mass, and triacylglycerol (TAG) accumulation. JHA treatments did not influence flight speed, but led to reductions in flight durations and flight distances. At day 3 posttreatment (PT), JHA‐treated females flew shorter times and less distance than the controls; JHA‐treated males, however, only flew shorter times than the controls. JHA treatments led to reductions in flight muscle mass in females at days 2–3 PT and reductions in TAG content in females at day 3 PT, but, these parameters were not influenced by JHA in males. These findings strongly support our hypothesis, from which we infer that JH is a major driver in C. medinalis oogenesis‐flight syndrome tradeoffs. Our data also reveal a JH‐sensitive stage in adulthood during which JH influences the oocyte‐flight syndrome in C. medinalis.