Flight Orientation Research Articles

Flight Orientation Behaviors Promote Optimal Migration Trajectories in High-Flying Insects

Many insects undertake long-range seasonal migrations to exploit temporary breeding sites hundreds or thousands of kilometers apart, but the behavioral adaptations that facilitate these movements remain largely unknown. Using entomological radar, we showed that the ability to select seasonally favorable, high-altitude winds is widespread in large day- and night-flying migrants and that insects adopt optimal flight headings that partially correct for crosswind drift, thus maximizing distances traveled. Trajectory analyses show that these behaviors increase migration distances by 40% and decrease the degree of drift from seasonally optimal directions. These flight behaviors match the sophistication of those seen in migrant birds and help explain how high-flying insects migrate successfully between seasonal habitats.

Release from bats: genetic distance and sensoribehavioural regression in the Pacific field cricket, Teleogryllus oceanicus

The auditory thresholds of the AN2 interneuron and the behavioural thresholds of the anti-bat flight-steering responses that this cell evokes are less sensitive in female Pacific field crickets that live where bats have never existed (Moorea) compared with individuals subjected to intense levels of bat predation (Australia). In contrast, the sensitivity of the auditory interneuron, ON1 which participates in the processing of both social signals and bat calls, and the thresholds for flight orientation to a model of the calling song of male crickets show few differences between the two populations. Genetic analyses confirm that the two populations are significantly distinct, and we conclude that the absence of bats has caused partial regression in the nervous control of a defensive behaviour in this insect. This study represents the first examination of natural evolutionary regression in the neural basis of a behaviour along a selection gradient within a single species.

What can be learnt from analysing insect orientation flights using probabilistic SLAM?

In this paper, we provide an analysis of orientation flights in bumblebees, employing a novel technique based on simultaneous localisation and mapping (SLAM) a probabilistic approach from autonomous robotics. We use SLAM to determine what bumblebees might learn about the locations of objects in the world through the arcing behaviours that are typical of these flights. Our results indicate that while the bees are clearly influenced by the presence of a conspicuous landmark, there is little evidence that they structure their flights to specifically learn about the position of the landmark.

Nesting Behavior of theTachysphex terminatusSpecies Group (Hymenoptera: Crabronidae)

Abstract The nesting behavior of species in the Tachysphex terminatus group is presented in phylogenetic order following Pulawski (1988). Eight taxa are treated: T. clarconis, T. antillarum, T. alpestris, T. linsleyi, T. terminatus, T. similis, T. a. apicalis, and T. apicalis fusus. This paper includes a literature review summary for each species, Howard E. Evans' unpublished field notes on T. linsleyi, T. terminatus, and T. similis, unreported insect museum prey records for T. terminatus and T. apicalis fusus, and long-term field studies on T. terminatus, T. similis, T. a. apicalis, and T. apicalis fusus. The field observations were made during the years 1960–2005 and include 929 nests excavated and examined at 52 locations in the central and eastern United States and southern Ontario. Emphasis is placed on: burrow excavation, tumulus leveling, temporary closure, orientation flight, hunting, prey capture, prey transport, nest entry, final closure, burrow design and length, cell depth, cell shape and size...

Environmental and pheromonal control of precopulatory behavior for synchronized mating in the white grub beetle, Dasylepida ishigakiensis (Coleoptera: Scarabaeidae)

Precopulatory behavior of the white grub beetle, Dasylepida ishigakiensis, was observed under laboratory conditions to determine the environmental factors controlling mating behavior, which is synchronized among individuals at dusk in the field. When light intensity was gradually decreased, both females and males sequentially started moving their antennae, legs and abdomen, then walking, and females rhythmically extruded their abdomen while males opened the elytra to fly. Pheromone release by females was suggested in a wind tunnel assay in which males oriented themselves to both intact and crushed bodies of females, but not of males. When illumination was suddenly turned off, precopulatory behavior of females and males was similarly induced but the frequencies of calling and flying behaviors were significantly reduced as compared with those observed when light intensity was gradually reduced. This result may suggest that gradual darkening is an important factor for their synchronized appearance from the soil and subsequent mating behaviors, including pheromone release in females and orientation flight in males. These behaviors were significantly suppressed at low temperatures below 18°C. This result supports our previous conclusion that temperature is the primary factor controlling the emergence and mating activity of sexually mature beetles in the field.

Effects of hive spacing, entrance orientation, and worker activity on nest relocation by honey bee queens

The mating flight is the riskiest period in the life of a honeybee queen. A major cause of queen mortality in apiaries may be the drifting of queens to foreign colonies. We investigated the effects of distance between hives, entrance orientation and worker activity on queen drifting. Only 4% of queens drifted in our experiments, all during their maiden orientation flight and all to the closest neighbouring hive. Neither drifting nor the length of time it took queens to relocate their hive was significantly affected by either entrance orientation or distance between hive stands (2 m or 5 m). However, queens took significantly longer to identify their hive and were more likely to drift when the number of workers at the entrance was lower than that at the neighbouring hive. Our results show that drifting can be low even when hives are placed in pairs with only 2 m between pairs, and that worker activity has an important role in guiding returning queens on their maiden orientation flight.

Running on uneven ground: leg adjustment to vertical steps and self-stability

Human running is characterized by comparably simple whole-body dynamics. These dynamics can be modelled with a point mass bouncing on a spring leg. Theoretical studies using such spring-mass models predict that running can be self-stable. In simulations, this self-stability allows for running on uneven ground without paying attention to the ground irregularities. Whether humans actually use this property of the mechanical system in such an irregular environment is, however, unclear. One way to approach this question is to study how the leg stiffness in stance and the leg orientation in flight are changed in response to ground perturbations. Here, for 11 human subjects we studied two consecutive contacts during running on uneven ground with a force plate of adjustable height (step of +5, +10 and +15 cm). We found that runners adjust their leg stiffness to the height of a vertical step. The adjustment is characterized by a 9% increase in leg stiffness in preparation for the perturbation and by a systematic decrease in proportion to the step height. At the highest vertical step (+15 cm), leg stiffness was reduced by about 26%. We also observed that the angle of attack decreased from 68 deg. to 62 deg. with increasing ground height. These leg adjustments are in accordance with the predictions of a stable spring-mass system. Furthermore, we could describe the identified leg forces and leg compressions with a simple spring-mass simulation for a given body mass, leg stiffness, angle of attack and initial conditions. We compared the experimental findings with the self-stabilizing properties of the spring-mass model, and discuss how humans use a combination of strategies that include purely mechanical self-stabilization and active neuromuscular control. Finally, beyond self-stability, we suggest that control may apply to smooth centre of mass kinematics.

Motion Processing Streams in Drosophila Are Behaviorally Specialized

Motion vision is an ancient faculty, critical to many animals in a range of ethological contexts, the underlying algorithms of which provide central insights into neural computation. However, how motion cues guide behavior is poorly understood, as the neural circuits that implement these computations are largely unknown in any organism. We develop a systematic, forward genetic approach using high-throughput, quantitative behavioral analyses to identify the neural substrates of motion vision in Drosophila in an unbiased fashion. We then delimit the behavioral contributions of both known and novel circuit elements. Contrary to expectation from previous studies, we find that orienting responses to motion are shaped by at least two neural pathways. These pathways are sensitive to different visual features, diverge immediately postsynaptic to photoreceptors, and are coupled to distinct behavioral outputs. Thus, behavioral responses to complex stimuli can rely on surprising neural specialization from even the earliest sensory processing stages.

Age and natural metabolically-intensive behavior affect oxidative stress and antioxidant mechanisms

Flying honey bees have among the highest mass-specific metabolic rates ever measured, suggesting that their flight muscles may experience high levels of oxidative stress during normal daily activities. We measured parameters of oxidative stress and antioxidant capacity in highly metabolic flight muscle and less active head tissue in cohorts of age-matched nurse bees, which rarely fly, and foragers, which fly several hours per a day. Naturally occurring foraging flight elicited an increase in flight muscle Hsp70 content in both young and old foragers; however catalase and total antioxidant capacity increased only in young flight muscle. Surprisingly, young nurse bees also showed a modest daily increase in Hsp70, catalase levels and antioxidant capacity, and these effects were likely due to collecting the young nurses soon after orientation flights. There were no differences in flight muscle carbonyl content over the course of daily activity and few differences in Hsp70, catalase, total antioxidant capacity and protein carbonyl levels in head tissue regardless of age or activity. In summary, honey bee flight likely produces high levels of reactive oxygen species in flight muscle that, when coupled with age-related decreases in antioxidant activity may be responsible for behavioral senescence and reduced longevity.

Two new PCR-RFLPs in the domestic pigeon (<i>Columba livia</i> var. <i>domestica</i>) lactate dehydrogenase A (<i>LDH-A</i>) gene (Brief report)

Abstract. Traditional selection of racing pigeons has been focusing on spatial orientation, velocity, and endurance of flight. LDHA gene is involved in aerobic and anaerobic metabolism of the muscle tissue (VAN HALL et al., 1999). Mutations in the LDHA gene can potentially diversify the homing performance of racing pigeons. Previously, two polymorphic sites of LDHA gene have been identified (DYBUS and KMIEĆ, 2002, DYBUS et al., 2006).

Temporal transition of nesting activities in the digger wasp Bembecinus tridens (Hymenoptera, Crabronidae)

A nest aggregation of Bembecinus tridens (Hymenoptera: Crabronidae) was investigated through observations of marked individual females in Northern Italy, in order to quantify frequencies, duration, and sequence of the nesting activities. Wasp females dug unicellular (in one case multicellular) nests preferably on oblique surfaces. Nest digging occurred mainly in the morning, while definitive nest closing was performed only in the afternoon, and nests were frequented by the owners for 3.52 days on average. One to eight nests were dug by single females in rapid succession in the area observed, revealing a rapid dispersion towards other sites. Provisioning and non‐provisioning (control) flights were positively correlated and occurred mainly in the late morning. A number of orientation flights followed most exits from the nests but was higher at the end of nest digging. Nest closing lasted more at the end of provisioning and at the end of digging. Provisioning began more frequently one day after nest digging (and oviposition), less frequently two days after, and the number of prey per day was negatively correlated with the number of provisioning days. Data are consistent with a situation of primitive progressive provisioning, and match that of other species of the same genus.

Visuo-motor learning during orientation flight of the fruit fly Drosophila

Visuo-motor learning during orientation flight of the fruit fly Drosophila

Synergism and redundancy in a plant volatile blend attracting grapevine moth females

A flight tunnel study was done to decipher the behavioral effect of grape odor in grapevine moth Lobesia botrana. A blend of 10 volatile compounds, which all elicit a strong antennal response, attracts mated grapevine moth females from a distance, by upwind orientation flight. These 10 grape volatiles are in part behaviorally redundant, since attraction to a 3-component blend of β-caryophyllene, ( E)-β-farnesene and ( E)-4,8-dimethyl-1,3,7-nonatriene was not significantly different from the 10-component blend. Blending these three compounds had a strong synergistic effect on female attraction, and omission of any one compound from this 3-component blend almost abolished attraction. It was nonetheless possible to substitute the three compounds with the other grape volatiles which are perceived by the female antenna, to partly restore attraction. Several blends, of varying composition, elicited significant attraction. The observed behavioral plasticity in response to grape volatile blends probably reflects the variation of the natural plant signal, since females oviposit on different grape varieties, in different phenological stages.

Appetitive flight patterns of male Agrotis segetum moths over landscape scales

An analysis is presented of the first harmonic radar studies of pheromone-plume locating flights of male Agrotis segetum moths over distances of up to 500 m. Upon release most moths flew in a direction having a downwind component. The first significant changes in flight orientations occur in the immediate vicinity of a pheromone source. Moths that were initially flying downwind change course and start flying crosswind whilst those that initially flew crosswind change course and start flying upwind. It is shown that such behaviour is consistent with the adoption of an effective plume-location strategy, and conditions are identified when downwind flights would be more advantageous than crosswind ones. Additionally, some of the complex flight patterns that can arise at later times are shown to be compatible with the adoption of an optimal biased scale-free (Lévy-flight) searching strategy. It is found that disruptive doses of sex pheromone can have a marked influence upon male moth flight patterns.

Spatial memory, navigation and dance behaviour in Apis mellifera

Navigation and dance communication in Apis mellifera have been extensively studied on the level of sensory processing, but the structure and content of the spatial memory underlying such phenomena have yet to be addressed. Here we survey new findings indicating that the memory used by bees to navigate within the range of their orientation flights is much more complex than hitherto thought. It appears to allow them to decide between at least two goals in the field, and to steer towards them over considerable distances. Two models concerning the structure of bees' spatial memory are developed from new empirical evidence. The first one relies on the integration of at least two flight vectors, while the second assumes the existence of a 'functional' map based on the information available on-site. These findings also raise questions about the process of encoding and decoding information in the context of the waggle dance. We review published data and recent evidence indicating that memories of topographical features might also be involved in dance communication, and point out what needs to be addressed to elucidate the corresponding memory demands. The flight paths of recruited bees can now be traced by means of radar techniques, and thus tools are available to tackle these questions.

Representation of behaviourally relevant information by blowfly motion-sensitive visual interneurons requires precise compensatory head movements

Flying blowflies shift their gaze by saccadic turns of body and head, keeping their gaze basically fixed between saccades. For the head, this results in almost pure translational optic flow between saccades, enabling visual interneurons in the fly motion pathway to extract information about translation of the animal and thereby about the spatial layout of the environment. There are noticeable differences between head and body movements during flight. Head saccades are faster and shorter than body saccades, and the head orientation is more stable between saccades than the body orientation. Here, we analyse the functional importance of these differences by probing visual interneurons of the blowfly motion pathway with optic flow based on either head movements or body movements, as recorded accurately with a magnetic search coil technique. We find that the precise head-body coordination is essential for the visual system to separate the translational from the rotational optic flow. If the head were tightly coupled to the body, the resulting optic flow would not contain the behaviourally important information on translation. Since it is difficult to resolve head orientation in many experimental paradigms, even when employing state-of-the-art digital video techniques, we introduce a 'headifying algorithm', which transforms the time-dependent body orientation in free flight into an estimate of head orientation. We show that application of this algorithm leads to an estimated head orientation between saccades that is sufficiently stable to enable recovering information on translation. The algorithm may therefore be of practical use when head orientation is needed but cannot be measured.

Orientation of Culex mosquitoes to carbon dioxide‐baited traps: flight manoeuvres and trapping efficiency

Females of Culex quinquefasciatus Say and Culex tarsalis Coquillet (Diptera: Culicidae) in the host-seeking stage were released and video recorded in three dimensions in a large field wind tunnel as they flew to four kinds of CO2-baited mosquito traps. The trapping efficiency (number of mosquitoes approaching compared to the number caught) was determined for each trap type. The Encephalitis Virus Surveillance (EVS), Mosquito Magnet Freedom (MMF) and Mosquito Magnet Liberty (MML) traps captured only 13-16% of approaching Cx. quinquefasciatus females, whereas the Mosquito Magnet-X (MMX) trap captured 58%. Similar results were obtained for Cx. tarsalis. Orientation behaviour and flight parameters of mosquitoes approaching the four traps were compared. Mosquitoes spent the most time orienting to the EVS trap. Flight speed decreased as mosquitoes entered the vicinity of each trap and a large portion of their time was spent within 30 cm downwind of the traps. Flights became highly tortuous downwind of the poorly performing traps and just upwind of the MMX trap. Differences between traps and possible explanations for the superior performance of the MMX trap are considered.

Cooperative random Lévy flight searches and the flight patterns of honeybees

The most efficient Lévy flight (scale-free) searching strategy for N independent searchers to adopt when target sites are randomly and sparsely distributed is identified. For N = 1 , it is well known that the optimal searching strategy is attained when μ = 2 , where the exponent μ characterizes the Lévy distribution, P ( l ) = l − μ , of flight-lengths. For N > 1 , the optimal searching strategy is attained as μ → 1 . It is suggested that the orientation flights of honeybees can be understood within the context of such an optimal cooperative random Lévy flight searching strategy. Upon returning to their hive after surveying a landscape honeybees can exchange information about the locations of target sites through the waggle dance. In accordance with observations it is predicted that the waggle dance can be disrupted without noticeable influence on a hive's ability to maintain weight when forage is plentiful.

Prior classical olfactory conditioning improves odour-cued flight orientation of honey bees in a wind tunnel

Odours are key cues used by the honey bee in various situations. They play an important role in sexual attraction, social behaviour and location of profitable food sources. Here, we were interested in the role of odours in orientation at short distance, for instance the approach flight to a floral patch or in close proximity to the hive entrance. Using a newly designed wind tunnel, we investigated the orientation behaviour of the bee towards two different odours: a social odour and a floral component, linalool. We then tested the effect of prior olfactory conditioning (conditioning of the proboscis extension reflex) on subsequent flight orientation. We showed that both stimuli induced orientated behaviour (orientated flights, circling around the odour source) in up to 70% of the worker bees, social odour being slightly more attractive than the linalool. We found thereafter that orientation performance towards the floral compound can be significantly enhanced by prior classical olfactory learning. This type of information transfer, from a Pavlovian associative context to an orientation task, might allow future foragers to acquire, within the hive, relevant information about the odours and food they will encounter during their later foraging bouts.

The use of waggle dance information by honey bees throughout their foraging careers

We studied the extent to which worker honey bees acquire information from waggle dances throughout their careers as foragers. Small groups of foragers were monitored from time of orientation flights to time of death and all in-hive behaviors relating to foraging were recorded. In the context of a novice forager finding her first food source, 60% of the bees relied, at least in part, on acquiring information from waggle dances (being recruited) rather than searching independently (scouting). In the context of an experienced forager whose foraging has been interrupted, 37% of the time the bees resumed foraging by following waggle dances (being reactivated) rather than examining the food source on their own (inspecting). And in the context of an experienced forager engaged in foraging, 17% of the time the bees initiated a foraging trip by following a waggle dance. Such dance following was observed much more often after an unsuccessful than after a successful foraging trip. Successful foragers often followed dances just briefly, perhaps to confirm that the kind of flowers they had been visiting were still yielding forage. Overall, waggle dance following for food discovery accounted for 12–25% of all interactions with dancers (9% by novice foragers and 3–16% by experienced foragers) whereas dance following for reactivation and confirmation accounted for the other 75–88% (26% for reactivation and 49–62% for confirmation). We conclude that foragers make extensive use of the waggle dance not only to start work at new, unfamiliar food sources but also to resume work at old, familiar food sources.