Flight activity of three Spodoptera spp., Spodoptera litura, S. exigua and S. depravata, measured by flight actograph

Flight activities of three Spodoptera species were measured by the aid of flight actograph: S. litura and S. exuiga being regarded as long‐distance migratory insects, and S. depravata being non‐migratory and diapause‐inducible species. In all species tested, flight activities were observed only in scotophase, males showed far higher activities than females, being several times higher at the time of maximum flight activity, which was observed within 2 days after adult eclosion. Total flight activity in males was highest in S. litura, some being flyable even 12 days after eclosion, followed by S. exigua being one‐third compared to the former species, while in S. depravata flight activity was nearly half of that of the second species and most ceased to fly within a week after eclosion. There occurred species‐specific daily rhythms in flight activity during respective scotophase. In S. litura, both females and males exhibited a peak of flight activity shortly after light‐off and exhibited the second flight activity in late scotophase, the females slightly but the males more actively compared to early scotophase. In S. exigua, both sexes did not respond to light‐off, did not show a peak of flight activity in early scotophase, whereas males, but not females prominently increased activity toward the end of scotophase. In S. depravata, both sexes exhibited a peak of flight activity in early scotophase, and the males revived flight activity, being maximum shortly before light‐on, but the females did not show a clear rhythm in flight activity. These features observed in flight activity were discussed in relation with migratory capability.     

Physical and physiological costs of ectoparasitic mites on host flight endurance

1. Dispersal is essential for locating mates, new resources, and to escape unfavourable conditions. Parasitism can impact a host's ability to perform energetically demanding activities such as long‐distance flight, with important consequences for gene flow and meta‐population dynamics. 2. Ectoparasites, in particular, can adversely affect host flight performance by diminishing flight aerodynamics and/or by inflicting physiological damage while feeding on host tissue. 3. Experimental flight assays were conducted using two fruit fly‐mite systems: Drosophila nigrospiracula (Patterson and Wheeler) – Macrocheles subbadius (Berlese) and D. hydei (Sturtevan) – M. muscaedomesticae (Scopoli). Flies that are burdened by mites are expected to exhibit lower flight endurance compared to uninfected flies. 4. The results show that the presence of mites (attached) significantly decreased flight endurance by 57% and 78% compared to uninfected D. nigrospiracula and D. hydei, respectively. The physiological damage caused by M. subbadius was revealed through a 53% decline in flight time among previously infected flies (mites removed just prior to flight assay). Surprisingly, the presumably phoretic M. muscaedomesticae also caused a 62% reduction in flight endurance among previously infected D. hydei. 5. These results suggest a strong deleterious effect of ectoparasitic mites on host flight performance, mediated by a reduction in flight aerodynamics and damage to host physiology. Adverse effects on host flight and/or dispersal may have broad implications for gene flow, population genetic structure, and local adaptation in both host and parasite meta‐populations.     

Experimental studies of migration in bugs of the genus Dysdercus

Summary Using a tethered flight technique, migration was studied in the African cotton stainer bugs Dysdercus fasciatus Sign., D. nigrofasciatus Stål, and D. superstitiosus (F.) (Heteroptera: Pyrrhocoridae). The flight system of males of all three species matures in 4–6 days post-eclosion, and flight can continue throughout life. Fed females of D. fasciatus do not fly, but rather histolyse the flight muscles and develop oocytes. Fed females of D. nigrofasciatus and D. superstitiosus exhibit some flight at 4 days post-eclosion, but flight is essentially absent at 6 days when flight muscle histolysis and oögenesis are in progress. Starved females of all three species undertake considerable flight and do not histolyse the flight muscles or develop oöcytes. In starved pairs coupling frequency is lowered permitting increased migration. The three species are thus facultative migrants using starvation as a releasing cue. D. fasciatus feeds on baobab (Adansonia) fruits which are seasonally abundant and is the most opportunistic species with the highest reproductive potential and no flight in the presence of food. D. nigrofasciatus feeds on a variety of herbaceous annuals and perennials; it follows an intermediate migration and reproductive strategy. Finally D. superstitiosus is the most catholic in its food habits and has the lowest reproductive potential. Both of the latter species retain some flight capacity even when fed, presumably so that individuals can locate suitable host plants.     

Intraspecific scaling of flight power in the bat Glossophaga soricina (Phyllostomidae)

Aerodynamic theory predicts that power output during flight should vary with body mass by an exponent of 1.56 when wing morphology remains constant (within an individual), and by an exponent of 1.19 when wing morphology changes with body mass (within a species or between species). I tested these predictions by estimating the power input during horizontal flight in three pregnant and two subadult Glossophaga soricina using a multivariate regression model. This analysis yielded power input during resting and flight as well as the energetic equivalent of change in body mass. A comparison of the estimated flight power for pregnant G. soricina, with published data on flight power of non-pregnant adults, revealed that energy turnover in flight is highest for pregnant G. soricina. Flight power of a 13-g pregnant G. soricina was even higher than that of a 16-g non-pregnant Glossophaga longirostris. A least-squares regression analysis yielded the following equations for the intraspecific scaling of flight power with body mass: power input during horizontal flight (P _f )=24099 body mass (bm; kg)^2.15 (r ²=0.97) for the intra-individual allometry (pregnancy) and P _f =113 bm(kg)^0.95 (r ²=0.99) for the inter-individual allometry (ontogeny). Both mass exponents are not significantly different from the predicted values for the scaling relationship of flight power within an individual (1.56) and within a species (1.19). This is the first measurement of power input during flight for subadult and pregnant bats. Accepted: 11 May 2000     

Flight initiations in Drosophila melanogaster are mediated by several distinct motor patterns

We have monitored the patterns of activation of five muscles during flight initiation of Drosophila melanogaster: the tergotrochanteral muscle (a mesothoracic leg extensor), dorsal longitudinal muscles #3, #4 and #6 (wing depressors), and dorsal ventral muscle #Ic (a wing elevator). Stimulation of a pair of large descending interneurons, the giant fibers, activates these muscles in a stereotypic pattern and is thought to evoke escape flight initiation. To investigate the role of the giant fibers in coordinating flight initiation, we have compared the patterns of muscle activation evoked by giant fiber stimulation with those during flight initiations executed voluntarily and evoked by visual and olfactory stimuli. Visually elicited flight initiations exhibit patterns of muscle activation indistinguishable from those evoked by giant fiber stimulation. Olfactory-induced flight initiations exhibit patterns of muscle activation similar to those during voluntary flight initiations. Yet only some benzaldehyde-induced and voluntary flight initiations exhibit patterns of muscle activation similar to those evoked by giant fiber stimulation. These results indicate that visually elicited flight initiations are coordinated by the giant fiber circuit. By contrast, the giant fiber circuit alone cannot account for the patterns of muscle activation observed during the majority of olfactory-induced and voluntary flight initiations.Abbreviations DLM/DLMn dorsal longitudinal muscle/motor neuron - DVM/DVMn dorsal ventral muscle/motor neuron - GF(s) giant fiber interneuron (s) - PSI peripherally synapsing interneuron - TTM/TTMn tergotrochanteral muscle/motor neuron     

The Influence of Food and Con-specifics on the Flight Potential of the Parasitoid <Emphasis Type="Italic">Ibalia leucospoides</Emphasis>

For insect parasitoids, knowledge of their flight capability is essential for a general understanding of the relationship with their hosts. For instance, flight capacity might partly determine their efficacy as biological control agents. Ibalia leucospoides Hochenwarch (Hymenoptera: Ibaliidae) is a solitary, pro-ovigenic parasitoid of the woodwasp, Sirex noctilio Boidin (Hymenoptera, Siricidae), an economically important pest of softwood forestation. This study explores the flight capacities of I. leucospoides females and assesses the effects of a sugar-rich food supply and crowding on female flight performance, by using computer-linked flight mills. The present study shows: (1) a high variability in flight potential of I. leucospoides females, (2) no effects of food supply and grouping on wasp flight (flight distance and speed), (3) a significant effect of body size and wing loading on flight performance and, (4) a significant body mass loss during the flight dependent on the total distance flown. The lack of effect of food on a highly energy-demanding activity as flight may be related to the life-history traits and nutritional strategies of this parasitoid. The relevance of these observations for the use of I. leucospoides as a biological control agent is discussed.     

Lifetime- and caste-specific changes in flight metabolic rate and muscle biochemistry of honeybees, Apis mellifera

Honeybees, Apis mellifera, who show temporal polyethism, begin their adult life performing tasks inside the hive (hive bees) and then switch to foraging when they are about 2–3 weeks old (foragers). Usually hive tasks require little or no flying, whereas foraging involves flying for several hours a day and carrying heavy loads of nectar and pollen. Flight muscles are particularly plastic organs that can respond to use and disuse, and accordingly it would be expected that adjustments in flight muscle metabolism occur throughout a bee’s life. We thus investigated changes in lifetime flight metabolic rate and flight muscle biochemistry of differently aged hive bees and of foragers with varying foraging experience. Rapid increases in flight metabolic rates early in life coincided with a switch in troponin T isoforms and increases in flight muscle maximal activities (V _max) of the enzymes citrate synthase, cytochrome c oxidase, hexokinase, phosphofructokinase, and pyruvate kinase. However, further increases in flight metabolic rate in experienced foragers occurred without additional changes in the in vitro V _max of these flight muscle metabolic enzymes. Estimates of in vivo flux (v) compared to maximum flux of each enzyme in vitro (fractional velocity, v/V _max) suggest that most enzymes operate at a higher fraction of V _max in mature foragers compared to young hive bees. Our results indicate that honeybees develop most of their flight muscle metabolic machinery early in life. Any further increases in flight metabolism with age or foraging experience are most likely achieved by operating metabolic enzymes closer to their maximal flux capacity.     

Effect of physiological factors on the flight ability of Calliptamus italicus L

Calliptamus italicus L. is a major migratory pest that often causes serious agricultural losses in the desert/semi‐desert steppe of Central Asia. The present study aims to understand the physiological factors that affect migration of C. italicus by examining the relationships between flight capacity, energy accumulation, ovarian development and Juvenile hormone (JH) titre. The results show that flight capacity decreases with age, being greatest among 2‐day‐old males and lowest among 13‐day‐old locusts. There is no significant correlation between energy accumulation and flight capacity, although JH titre is negatively correlated with flight capacity. Energy accumulation and flight capacity first increases then decreases in 1–13‐day‐old females. Glycogen accumulation is significantly correlated with flight speed, and triglyceride accumulation is significantly correlated with flight distance, duration and speed. Changes in JH titre in 1–13‐day‐old females show double titre peaks, which are negatively correlated with flight capacity. Seven‐day‐old C. italicus have the highest glycogen and triglyceride accumulation, greatest flight capacity, grade II ovarian development and lowest JH titre. By contrast, 13‐day‐old C. italicus have the lowest triglyceride accumulation, lowest flight capacity, grade IV ovarian development and lowest JH titre. Taken together, these findings demonstrate that there is a trade‐off between ovarian development and flight capacity of C. italicus.     

The relationship between pre‐oviposition flight behaviour and reproductive timing in whitefly parasitoids

Although the timing of oögenesis appears to be a major life‐history organizer in the parasitoid Hymenoptera, relatively little is known about how this trait correlates with population dispersal by flight in these wasps. Pre‐oviposition flight behaviours of 1‐day‐old female wasps in a vertical flight chamber are measured to test correlations between these traits and with reproductive timing. The focus of the present study is on two genera of whitefly parasitoids (Encarsia and Eretmocerus) that differ in reproductive timing when feeding on a shared host (Bemisia tabaci). The two Eretmocerus species engage in vertical flight within 3 min of release far more frequently than the three Encarsia species. Because the former typically possess a more time‐limited reproductive strategy than the latter, this lends support for a positive interspecific association between early‐life reproduction and early‐life flight incidence. Within species, however, egg load does not correlate with flight propensity for any of the tested species. Furthermore, in Eretmocerus eremicus, the relationship between effective flight distance (i.e. the product of vertical climb rate and flight duration) and egg load appears to be labile rather than fixed because different trials show evidence for either negative or positive correlations between these variables. The source of this context dependence may be variation in either biotic (e.g. longevity) or abiotic (e.g. temperature and relative humidity) factors.     

Comparison of dispersal ability between the green leafhoppers,Nephotettix virescens andN. cincticeps (hemiptera: Cicadellidae) by tethered flight

Dispersal propensity of green leafhoppers was compared between a tropical species, Nephotettix virescens, and a temperate species, N. cincticeps. The flight ability was measured with tethered flight technique under laboratory conditions (25°C, 16L-8D). The pre-flight period was shorter and the flight duration was longer in N. virescens than in N. cincticeps in both sexes. No significant correlations were found between the flight activity and morphometric characters for either of the two species. The results suggested that dispersal propensity of N. virescens is higher than that of N. cincticeps.     

Time‐restricted flight ability influences dispersal and colonization rates in a group of freshwater beetles

Variation in the ability to fly or not is a key mechanism for differences in local species occurrences. It is increasingly acknowledged that physiological or behavioral mechanisms rather than morphological differences may drive flight abilities. However, our knowledge on the seasonal variability and stressors creating nonmorphological differences in flight abilities and how it scales to local and regional occurrences is very limited particularly for small, short‐lived species such as insects. Here, we examine how flight ability might vary across seasons and between two closely related genera of freshwater beetles with similar geographical ranges, life histories, and dispersal‐related morphology. By combining flight experiments of >1,100 specimens with colonization rates in a metacommunity of 54 ponds in northern and eastern Europe, we have analyzed the relationship between flight ability and spatio‐environmental distribution of the study genera. We find profound differences in flight ability between the two study genera across seasons. High flight ability for Acilius (97% of the tested individuals flew during the experiments) and low for Graphoderus (14%) corresponded to the different colonization rates of newly created ponds. Within a 5‐year period, 81 and 31% of the study ponds were colonized by Acilius and Graphoderus, respectively. While Acilius dispersed throughout the season, flight activity in Graphoderus was restricted to stressed situations immediately after the emergence of adults. Regional colonization ability of Acilius was independent of spatial connectivity and mass effect from propagule sources. In contrast, Graphoderus species were closely related to high connectivity between ponds in the landscape. Our data suggest that different dispersal potential can account for different local occurrences of Acilius and Graphoderus. In general, our findings provide some of the first insights into the understanding of seasonal restrictions in flight patterns of aquatic beetles and their consequences for species distributions.     

Quantitative studies on the release of locust adipokinetic hormone

Fractionation of methanolic extracts of haemolymph on Sephadex LH-20 made possible the measurement of the titre of adipokinetic hormone in the haemolymph of locusts. Experimentally produced high concentrations of haemolymph carbohydrate caused a delay in the mobilization of lipid during flight, and very low titres of the hormone were present in the haemolymph of locusts injected with trehalose immediately before a 25 min flight. In these locusts flight speed was higher than saline-injected controls. Although delayed lipid mobilization during flight was also seen in locusts injected with sucrose, sucrose is not utilized for flight metabolism and flight speed was not increased by the injection. Tentative estimates of the release rate (c. 1000pg/20min flight) and half life (c. 20 min) of adipokinetic hormone during flight are made. The results described suggest that during flight the rate at which trehalose disappears from the haemolymph does not play a major role in the initiation of the release of adipokinetic hormone.     

Exhaled air temperature as a function of ambient temperature in flying and resting ducks

Exhaled air temperature (T _exh) has a paramount effect on respiratory water loss during flight. For migratory birds, low T _exh potentially reduces water loss and increases flight range. However, only three studies provide empirical data on T _exh during flight. The aim of this study was to record T _exh of birds during rest and flight at a range of controlled ambient temperatures (T _amb). One wigeon and two teal flew a total of 20 times in a wind tunnel at T _amb ranging from 1° to 24°C. T _exh during flight did not differ between the two species and was strongly correlated with T _amb (T _exh=1.036 T _amb + 13.426; R²=0.58). In addition, body temperature had a weak positive effect on T _exh. At a given T _amb, T _exh was about 5°C higher during flight than at rest.     

Flight Speed of Tethered Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae) Alates

Alates of the Eastern subterranean termite, Reticulitermes flavipes (Kollar) were collected over two flight seasons (2002 and 2004) and flown on flight mills. Data were collected to test if alate mass, colony origin, or gender influenced flight speed. Flight speed ranged from 3.14 to 69.12 cm s⁻¹ and the maximum distance flown by an alate was 458.3 m. Alate mass (P = 0.9406), gender (P = 0.3976), colony origin (P = 0.1244), and the interaction of gender and colony (P = 0.7093) did not significantly influence flight speed. Additionally, an electronic counting device was used to provide instantaneous flight speeds and allowed flight speed to be modeled during acceleration, cruising, and deceleration periods of flight. Mean (±SEM) flight speeds in 2004 were 20.64 (±2.21) cm s⁻¹ (n = 13) for males and 17.76 cm s⁻¹ (n = 1) for the single female flown, falling within the range of the 2002 values.     

Neuromodulation of flight initiation by octopamine in the cockroach Periplaneta americana

We have tested the effect of a known insect neuromodulator, octopamine, on flight initiation in the cockroach. Using minimally dissected animals, we found that octopamine lowered the threshold for windevoked initiation of flight when applied to either of two major synaptic sites in the flight circuitry: 1) the last abdominal ganglion, where wind-sensitive neurons from the cerci excite dorsal giant interneurons, or 2) the metathoracic ganglion, where the dorsal giant interneurons activate interneurons and motoneurons which are involved in producing the rhythmic flight motor pattern in the flight muscles (Fig. 2).Correlated with this change in flight initiation threshold, we found that octopamine applied to the last abdominal ganglion increased the number of action potentials produced by individual dorsal giant interneurons when recruiting the cereal wind-sensitive neurons with wind puffs (Figs. 3, 4, 5) or with extracellular stimulation of their axons (Fig. 6). Octopamine increases the excitability of the giant interneurons (Figs. 7, 8). Also, when we stimulated individual dorsal giant interneurons intracellularly, the number of action potentials needed to initiate flight was reduced when octopamine was applied to the metathoracic ganglion (Fig. 9).Abbreviations EMG electromyogram - dGIs dorsal giant interneurons - GI giant interneuron - A6 sixth abdominal ganglion - T3 third thoracic ganglion - EPSP excitatory postsynaptic potential     

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.     

Flight Behaviour of Merolimnic Insects from the Leutra River (Thuringia,Germany)

1. The flight behaviour of adult merolimnic insects was studied on the first order limestone stream Leutra (Thuringia, Germany) using sticky traps. The focus of the present study was on testing the colonisation cycle hypothesis of Müller (1954) and on the small-scale dispersal of adult merolimnic insects. 2. A high number of sticky traps was used to guarantee a proper statistical analysis of the data and exclude effects of the heterogeneous environment on flight behaviour. 3. The flight behaviour of Leuctra hippopus, Nemoura cambrica, Nemoura flexuosa, Protonemura nitida, Protonemura praecox (Plecoptera), and Baetis rhodani (Ephemeroptera) was studied. 4. Specific flight corridors were observed for P. praecox, Nemoura spp., L. hippopus and B. rhodani. 5. Comparison of the catches on sticky traps set perpendicular to the water showed no significant flight upstream in any of the taxa studied. Thus, the results do not support the hypothesis of the colonisation cycle.     

Molecular and functional characterization of the flightin gene in the Oriental fruit fly,Bactrocera dorsalis

Flightin is a protein in flight muscles and is crucial for the flight capacity. Flightin also has been proposed as a protein with deep ancestry and functions outside of flight muscles. However, functional and molecular characterization of flightin achieved so far is mainly in flight muscles of Drosophila. Here, we cloned the flightin (Bd-flightin) gene and tested its expression and function in Bactrocera dorsalis, an important migratory pest. Phylogenetic analysis based on flightin orthologs revealed that the divergence of flightin is consistent with the taxonomic classification of insects. Motif analysis indicated obvious variations in flightin orthologs, which may have occurred during speciation and functional differentiation. The expression is quite low during egg and larval stages, which largely increased during pupal stage and then peaked at the beginning of the adult stage. Bd-flightin also showed tissue- and age-specific expression patterns during adult stage. The relative expression level is low in wing, head, ovary and testis, which is relatively higher in leg and abdominal wall and much higher in thorax. Injection of late pupae and newly eclosed adults with 1 μg flightin dsRNA per insect both significantly reduced the expression of flightin and the flight capacity in males and females. In addition, silencing the expression of flightin also decreased the weight ratio of thorax and whole-body. These results suggested that flightin plays important roles in flight muscle development and flight function in B. dorsalis, which can potentially be used to control the flight behaviour of the fruit fly.     

Flight potential of <Emphasis Type="Italic">Microplitis mediator</Emphasis>, a parasitoid of various lepidopteran pests

The efficacy of using natural enemies to control pests under field conditions largely depends on their mobility and, more specifically, on their capacity to quickly locate pest infestation. For many natural enemies, for example parasitoids, mobility is directly related to flight aptitude, which is determined by the capacity and inclination of a species to engage in flight. In this study, we determined the various factors that affected flight performance of Microplitis mediator (Haliday) (Hymenoptera: Braconidae), using a computer-monitored flight mill. No differences were found in flight performance (i.e., flight distance, duration, speed) between both sexes of M. mediator, and flight capacity increased up to an age of 5–7 days followed by a gradual decline afterwards. For one-day-old female parasitoids, mean (±SE) flight distance and duration were 6.23 ± 0.88 km and 85.15 ± 14.44 min, respectively, with a maximum flight distance of 18.0 km. For male parasitoids, mean flight distance and duration were 5.27 ± 0.51 km and 85.74 ± 7.63 min, respectively. Mating status did not affect flight performance of males, while flight distance of four-day-old ovipositing M. mediator females was much lower than that of un-mated females of the same age. Un-mated adults flew most actively at 22–24°C and inclination to fly gradually declined with decreasing temperature. Temperatures above 26°C considerably reduced flight activity of M. mediator. Wasps engaged in normal flight under a broad range of relative humidity (RH) conditions, with an optimum RH range identified as 75–90%. Our research shows that M. mediator is a highly active parasitoid, because both sexes show great inclination to fly under a range of environmental conditions and flight capacity at different ages. Our results can help explain parasitoid performance in the field and provide baseline information to help guide augmentative releases. Handling editor: Dirk Babendreier.