Time-correlated flights of juvenile and mature locusts: A comparison between free and tethered animals

The flights of free and tethered Locusta migratoria were followed from initiation with a high-speed film camera. A longer sequence of wing-beat cycles can thus be correlated unequivocally with the animals's movement in time and space. In both flight situations the locusts start with approximately the same instantaneous wing-beat frequency. During the early flight phase free-flying animals increase their wing-beat frequency, whereas for tethered locusts this parameter remains constant or even decreases. The general flight pattern is similar in juvenile and mature locusts; the juveniles however, fly with alower wing-beat frequency and flight speed. The differences in the wing-beat frequencies for both flight performances are discussed with respect to differences in the sensory inputs to the flight motor centre.     

An ultrastructural study of tettigoniid muscle in relation to function

The modian dorsal longitudinal indirect flight muscles from the mesothorax and metathorax of Homorocoryphus nitidulus vicinus have been studied to determine whether structural differences might offer an explanation for reports that the mesothoracic musculature effects a wing-beat rate of 140 beats/sec during stridulation, whereas during flight, it, like that of the metathorax, effects wing-beat frequencies of 14 to 20 beats/sec. No differences were observed and it is concluded that the high wing-beat rate, reported during stridulation, is not reflected in any specific modification of mesothoracic muscle fine structure.     

Post-larval development of two rhythmical behavioural patterns: flight and song in the grasshopper,Omocestus viridulusS*

The ontogeny of two rhythmical outputs, wing-beat in flight and leg stroke in stridulation, was followed during post-larval development of the grasshopper 0.viridulus, The wing-beat frequency increases by a factor of c. 2; the repetition rate of the leg strokes remains constant. These results are discussed with respect to a hypothesis of neuronal economy within the CNS, and their biological significance.     

Mitochondria in the flight muscles of insects. III. Mitochondrial cytochrome c in relation to the aging and wing beat frequency of flies

1. In the present study a correlation has been sought between aging, flight muscle mitochrondria (sarcosomes), cytochrome c, and flight ability in the blowfly, Phormia regina. 2. During the 1st week of adult life, individual sarcosomes increase in mass from 2.7 x 10^–7 µg. dry weight at the time of emergence, to 8.5 x 10^–7 µg. by the 7th day. During this period of growth, the number of sarcosomes per fly (6.7 x 10⁸) remains constant. When mature, the sarcosomes account for 32.6 per cent of the total muscle dry weight, or close to 40 per cent on a wet weight basis. 3. It appears probable that the high content of flight muscle cytochromes is entirely localized in the sarcosomes. The cytochromes continue to be synthesized and increase in titer within the sarcosomes for 7 days after adult emergence. 4. As determined spectroscopically, the various cytochrome components at all times maintain a constant ratio both to one another and to the sarcosomal dry weight. This suggests the possibility that the cytochrome system may be synthesized as a single entity. 5. The wing-beat frequency of Drosophila funebris and Phormia varies with the age of these flies, being lowest at the time of emergence and maximum after the 6th day. 6. The relations between wing-beat frequency, respiration during flight, and sarcosomal cytochrome c content are discussed. On the basis of some likely assumptions it is calculated that the cytrochrome c turnover number is over 5,000, and that the cytochrome c turns over once for every two wing-beat cycles.     

Effects of ambient temperature,humidity, and age on wing-beat frequency of Periplaneta species

Wing-beat frequency of 2-week-old male Periplaneta americana cockroaches was measured during tethered flight at ambient temperatures from 19 to 35°C and 50 and 95% r.h. Between 19 and 27°C the frequency increased at a nearly constant rate (0·6 Hz/deg at 50% r.h.; 0·7 Hz/deg at 95% r.h.), but the rate of increase decreased markedly between 27 and 35°C. This decline differed from results of past investigations of wing-beat frequency for this species. Wing-beat frequencies at 95% r.h. were significantly higher than those at 50% r.h. for ambient temperatures between 27 and 35°C. reductions in wing-beat frequency at high temperatures and differences in wing-beat frequency between 50 and 95% r.h. may reflect decreases in internal thoracic temperature resulting from cuticular water loss.     

The influence of age and culture-temperature on the wing-beat frequency of the migratory locust, Locusta migratoria

Starting on the day of adult ecdysis, the wing-beat frequency of Locusta migratoria rises exponentially. The initial frequency for both sexes is ca. 10 Hz, and the final frequency for males (24–25 Hz) is somewhat higher than for females (ca. 21 Hz). Initial and final frequencies appear to be genetically programmed, because they are independent of the culture cage temperature and amount of flight experience. The rate at which the final frequency is attained, however, is culture temperature dependent, being faster when the temperature is higher. After having reached the final frequency the wing-beat changes only a little up to the time of the animal's death.     

Flying in tune: sexual recognition in mosquitoes

Mosquitoes hear with their antennae, which in most species are sexually dimorphic. Johnston, who discovered the mosquito auditory organ at the base of the antenna 150 years ago, speculated that audition was involved with mating behaviour. Indeed, male mosquitoes are attracted to female flight tones. The male auditory organ has been proposed to act as an acoustic filter for female flight tones, but female auditory behavior is unknown. We show, for the first time, interactive auditory behavior between males and females that leads to sexual recognition. Individual males and females both respond to pure tones by altering wing-beat frequency. Behavioral auditory tuning curves, based on minimum threshold sound levels that elicit a change in wing-beat frequency to pure tones, are sharper than the mechanical tuning of the antennae, with males being more sensitive than females. We flew opposite-sex pairs of tethered Toxorhynchites brevipalpis and found that each mosquito alters its wing-beat frequency in response to the flight tone of the other, so that within seconds their flight-tone frequencies are closely matched, if not completely synchronized. The flight tones of same-sex pairs may converge in frequency but eventually diverge dramatically.     

Genetic Variability of Flight Metabolism in DROSOPHILA MELANOGASTER. I. Characterization of Power Output during Tethered Flight

The mechanical power imparted to the wings during tethered flight of Drosophila melanogaster is estimated from wing-beat frequency, wing-stroke amplitude and various aspects of wing morphology by applying the steady-state aerodynamics model of insect flight developed by Weis-Fogh (1972, 1973). Wing-beat frequency, the major determinant of power output, is highly correlated with the rate of oxygen consumption. Estimates of power generated during flight should closely reflect rates of ATP production in the flight muscles, since flies do not acquire an oxygen debt or accumulate ATP during flight. In an experiment using 21 chromosome 2 substitution lines, lines were a significant source of variation for all flight parameters measured. Broadsense heritabilities ranged from 0.16 for wing-stroke amplitude to 0.44 for inertial power. The variation among lines is not explained by variation in total body size (i.e., live weight). Line differences in flight parameters are robust with respect to age, ambient temperature and duration of flight. These results indicate that characterization of the power output during tethered flight will provide a sensitive experimental system for detecting the physiological effects of variation in the structure or quantity of the enzymes involved in flight metabolism.     

Estimating the relative fitness of local adaptive peaks: the aerodynamic costs of flight in mimetic passion-vine butterflies Heliconius

In a related paper, we demonstrated that mimetic Heliconius butterflies have converged in wing-beat frequency and degree of asymmetry in the wing motion, whereas sister species are dissimilar in these same traits. Warning signals of sympatric, distasteful species converge in evolutionary models in order to educate their predators more efficiently that the signal is associated with unprofitable prey. Barring other constraints, the behaviours of the different co-mimetic pairs should ultimately converge on that behaviour which minimizes the energetic cost of flight. We estimated the energetic cost of each mimic''s flight behaviour in order to predict the difference in height of each fitness peak and the direction of convergent selection qualitatively. Following adjustments for body mass, mimetic Heliconius melpomene and Heliconius erato required more aerodynamic power than Heliconius cydno and Heliconius sapho. This difference was attributed to the slower flight speeds and higher wing-beat frequencies of H. melpomene and H. erato. Consequently, H. melpomene and H. erato expended more energy per unit distance per unit body mass than H. cydno and H. sapho. However, differences in body mass may equalize energy budgets and stabilize the sympatric coexistence of the two pairs of co-mimics.     

Effects of temperature on properties of flight neurons in the locust

High ambient temperatures increase the wing-beat frequency in flying locusts, Locusta migratoria. We investigated parameters of circuit and cellular properties of flight motoneurons at temperatures permissive for flight (20–40 °C). As the thoracic temperature increased motoneuronal conduction velocity increased from an average of 4.40 m/s at 25 °C to 6.73 m/s at 35 °C, and the membrane time constant decreased from 11.45 ms to 7.52 ms. These property changes may increase locust wing-beat frequency by affecting the temporal summation of inputs to flight neurons in the central circuitry. Increases in thoracic temperature from 25–35 °C also resulted in a hyperpolarization of the resting membrane potentials of flight motoneurons from an average of-41.1 mV to -47.5 mV, and a decrease of input resistances from an average of 3.45 M to 2.00 M. Temperature affected the measured input resistance both by affecting membrane properties, and by altering synaptic input. We suggest that the increase in conduction velocity Q₁₀=1.53) and the decrease of membrane time constant (Q₁₀=0.62) would more than account for the wing-beat frequency increase (Q₁₀=1.15). Hyperpolarization of the resting membrane potential (Q₁₀=1.18) and reduction in input resistance (Q₁₀=0.54) may be involved in automatic compensation of temperature effects.Abbreviations ANOVA analysis of variance - CPG central pattern generator - DL dorsal longitudinal muscles - EMG electromyographic - MN motoneuron - PSP post synaptic potential - Q₁₀ temperature coefficient - RMP resting membrane potential - S.D. standard deviation - SR stretch receptor     

Alternative versions of the myosin relay domain differentially respond to load to influence Drosophila muscle kinetics

We measured the influence of alternative versions of the Drosophila melanogaster myosin heavy chain relay domain on muscle mechanical properties. We exchanged relay domain regions (encoded by alternative versions of exon 9) between an embryonic (EMB) isoform and the indirect flight muscle isoform (IFI) of myosin. Previously, we observed no effect of exchanging the EMB relay domain region into the flight muscle isoform (IFI-9b) on in vitro actin motility velocity or solution ATPase measurements compared to IFI. However, in indirect flight muscle fibers, IFI-9b exhibited decreased maximum power generation (P_max) and optimal frequency of power generation (f_max) to 70% and 83% of IFI fiber values. The decrease in muscle performance reduced the flight ability and wing-beat frequency of IFI-9b Drosophila compared to IFI Drosophila. Previously, we found that exchanging the flight muscle specific relay domain into the EMB isoform (EMB-9a) prevented actin movement in the in vitro motility assay compared to EMB, which does support actin movement. However, in indirect flight muscle fibers EMB-9a was a highly effective motor, increasing P_max and f_max 2.5-fold and 1.4-fold, respectively, compared to fibers expressing EMB. We propose that the oscillatory load EMB-9a experiences in the muscle fiber reduces a high activation energy barrier between two strongly bound states of the cross-bridge cycle, thereby promoting cross-bridge cycling. The IFI relay domain's enhanced sensitivity to load increases cross-bridge kinetics, whereas the EMB version is less load-sensitive.     

A comparative study of flight performance and fuel utilization as a function of age in females of Florida mosquitoes

Sugar-fed females of 7 species of Florida mosquitoes were flown for 4·5 hr on a flight-mill system twice a week during their life span and analysed for flight performance (speed of flight during the first hour, the distance flown at the end of 4·5 hr flight period, and per cent reduction in flight speed of the fourth hour as compared to the first hour) and for utilization of haemolymph sugars, glycogen, and triglycerides reserves.Species-specific differences occurred in 50 per cent survival age and flight performance. Aedes taeniorhynchus, Mansonia titillans, Culex nigripalpus, Psorophora confinnis, and A. sollicitans exhibited maximum flight potential during 2 to 8 weeks after emergence, as compared with only 1 to 2 weeks in A. aegypti and Anopheles quadrimaculatus. This suggested that the former group of mosquitoes possessed a greater potential for dispersal by searching flights during their life span when compared with the latter group. Glycogen was the only energy reserve utilized during sustained tethered flight in all species. The energy requirements as calculated from depletion of glycogen during sustained flight for 4·5 hr in different species varied from 0·06 to 0·09 cal/1000 m or 16 to 32 cal/hr per g. The considerably lower values of energy expended, on the average of 16 to 22 cal/hr per g in C. nigripalpus, P. confinnis, A. sollicitans, and A. aegypti, compared with 30 to 32 cal/hr per g in A. taeniorhynchus and M. titillans, is most probably due to the substantially lower speed of the former group of mosquitoes, 1000 to 1500 m/hr, than in the latter group of mosquitoes, 1500 to 2000 m/hr. A. sollicitans and M. titillans started to show wing abrasion and distinct signs of senescence during the last 2 to 3 weeks of their life span.Non-flown females of different species maintained on sugar ab lib. achieved maximum levels of glycogen and triglycerides reserves during the second week and they maintained these levels for the major portion of their life span, after which the levels of these reserves showed a distinct decline. It is suggested that this stabilization and decline of the energy reserves control longevity and flight potential in each mosquito species.     

Alternative S2 Hinge Regions of the Myosin Rod Affect Myofibrillar Structure and Myosin Kinetics

The subfragment 2/light meromyosin “hinge” region has been proposed to significantly contribute to muscle contraction force and/or speed. Transgenic replacement of the endogenous fast muscle isovariant hinge A (exon 15a) in Drosophila melanogaster indirect flight muscle with the slow muscle hinge B (exon 15b) allows examination of the structural and functional changes when only this region of the myosin molecule is different. Hinge B was previously shown to increase myosin rod length, increase A-band and sarcomere length, and decrease flight performance compared to hinge A. We applied additional measures to these transgenic lines to further evaluate the consequences of modifying this hinge region. Structurally, the longer A-band and sarcomere lengths found in the hinge B myofibrils appear to be due to the longitudinal addition of myosin heads. Functionally, hinge B, although a significant distance from the myosin catalytic domain, alters myosin kinetics in a manner consistent with this region increasing myosin rod length. These structural and functional changes combine to decrease whole fly wing-beat frequency and flight performance. Our results indicate that this hinge region plays an important role in determining myosin kinetics and in regulating thick and thin filament lengths as well as sarcomere length.     

Comparative characteristics of the wing apparatus and flight of Syrphid flies (Diptera: Syrphidae)

Comparative analysis of the wing apparatus and flight in nine species of flower flies (Syrphidae) has been performed. Data on the flight velocity, aerodynamic force, wing-beat frequency, stroke amplitude and stroke plane angle, wing area, body mass and volume, as well as correlations between these parameters at the intraspecific and family levels, have been obtained. Based on the obtained data, the subfamilies Syrphinae and Eristalinae have been compared.     

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.     

FORELIMB POSTURE IN DINOSAURS AND THE EVOLUTION OF THE AVIAN FLAPPING FLIGHT-STROKE

Ontogenetic and behavioral studies using birds currently do not document the early evolution of flight because birds (including juveniles) used in such studies employ forelimb oscillation frequencies over 10 Hz, forelimb stroke-angles in excess of 130°, and possess uniquely avian flight musculatures. Living birds are an advanced morphological stage in the development of flapping flight. To gain insight into the early stages of flight evolution (i.e., prebird), in the absence of a living analogue, a new approach using Strouhal number     was used. Strouhal number is a nondimensional number that describes the relationship between wing-stroke amplitude ( A ), wing-beat frequency ( f ), and flight speed ( U ). Calculations indicated that even moderate wing movements are enough to generate rudimentary thrust and that a propulsive flapping flight-stroke could have evolved via gradual incremental changes in wing movement and wing morphology. More fundamental to the origin of the avian flapping flight-stroke is the question of how a symmetrical forelimb posture—required for gliding and flapping flight—evolved from an alternating forelimb motion, evident in all extant bipeds when running except birds.     

Swarming behaviour of the mosquito Culex pipiens quinquefasciatus: a quantitative analysis

ABSTRACT. Analysis of video recordings of Culex pipiens quinquefasciatus Say swarms showed that the overall swarming patterns of males and virgin females are similar, even though the short-term characteristics of their flight, such as speed and turning angle, differ. This suggests they have the same response to the visual cue of the swarm 'marker'. The swarming pattern of an individual consists of elliptical loops which, in the short-term, define an area smaller than the swarm as a whole. The foci of these ellipses drift gradually with respect to the marker; individuals do not seem to have preferred positions within the swarm, but drift at random. Male mosquitoes identify and locate females by the sound of their wing-beats. Diffuse sound at the wing-beat frequency of female C.p. quinquefasciatus (500–600 Hz) caused males within the swarm to slow down significantly. Their turning behaviour remained unchanged, so the ellipses decreased in size. The swarm as a whole collapsed into a smaller volume, centred over the marker. Apparently, the response of males to sound consists of at least two parts: initially they alter their flight speed, and then alter their turning behaviour once they have located the source of the sound.     

Do Levant Sparrowhawks Accipiter brevipes also migrate at night?

Flight paths of visually identified Levant Sparrowhawks Accipiter brevipes on autumn migration were analysed with a tracking radar in the Arava Valley, Israel. This time of the year there are no significant numbers of other species with a similar wing-beat pattern. This wing-beat pattern was found not only in daytime but also frequently at night. It is suggested that the Levant Sparrowhawk uses two strategies of migration: (1) soaring and gliding to reduce energy consumption; (2) flapping flight to reduce time spent on migration. The latter may be more important towards the end of the migratory season and/or when birds have become separated from the main migratory stream.     

Carbohydrate and fatty acid titres during flight of the migrant noctuid moth,Anticarsia gemmatalis Hübner

Haemolymph concentrations of total carbohydrate and fatty acids were determined in velvetbean caterpillar (Anticarsia gemmatalis Hübner, Lepidoptera: Noctuidae) adult females throughout a 4-hr period of tethered flight. Total carbohydrate concentration decreased from approx. 30 to 10 μg/μl during the first 45 min of flight. Total fatty acid concentration increased from approx. 20 to 40 μg/μl during the first 60 min of flight and then declined to and stabilized at preflight levels. The decrease in wet weight (from approx. 97 to 80 mg/moth) during flight was probably due to defecation since no change in dry weight or haemolymph volume occurred. After 4 hr of flight, no apparent change in whole body lipid content (approx. 12 mg/moth) was observed but the much smaller carbohydrate content was reduced approx. 80% (from approx. 0.6 to 0.1 mg/moth). Approximately equal amounts (approx. 360–550 μg) of carbohydrate and lipid were removed from the haemolymph during 4 hr of flight. Changes in the haemolymph concentrations of palmitic, oleic and linoleic acids correspond to the changes in total fatty acid concentration of the haemolymph, indicating that these are the major components of the lipid mobilized and utilized during flight of A. gemmatalis.     

Triglyceride pools, flight and activity variation at the Gpdh locus in Drosophila melanogaster

We have created a set of P-element excision-derived Gpdh alleles that generate a range of GPDH activity phenotypes ranging from zero to full activity. By placing these synthetic alleles in isogenic backgrounds, we characterize the effects of minor and major activity variation on two different aspects of Gpdh function: the standing triglyceride pool and glycerol-3-phosphate shuttle-assisted flight. We observe small but statistically significant reductions in triglyceride content for adult Gpdh genotypes possessing 33-80% reductions from normal activity. These small differences scale to a notable proportion of the observed genetic variation in triglyceride content in natural populations. Using a tethered fly assay to assess flight metabolism, we observed that genotypes with 100 and 66% activity exhibited no significant difference in wing-beat frequency (WBF), while activity reductions from 60 to 10% showed statistically significant reductions of approximately 7% in WBF. These studies show that the molecular polymorphism associated with GPDH activity could be maintained in natural populations by selection in the triglyceride pool.