Visual ground pattern modulates flight speed of male Oriental fruit moth Grapholita molesta

Insects flying in a horizontal pheromone plume must attend to visual cues to ensure that they make upwind progress. Moreover, it is suggested that flying insects will also modulate their flight speed to maintain a constant retinal angular velocity of terrestrial contrast elements. Evidence from flies and honeybees supports such a hypothesis, although tests with male moths and beetles flying in pheromone plumes are not conclusive. These insects typically fly faster at higher elevations above a high‐contrast ground pattern, as predicted by the hypothesis, although the increase in speed is not sufficient to demonstrate quantitatively that they maintain constant visual angular velocity of the ground pattern. To test this hypothesis more rigorously, the flight speed of male oriental fruit moths (OFM) Grapholita molesta Busck (Lepidoptera: Tortricidae) flying in a sex pheromone plume in a laboratory wind tunnel is measured at various heights (5–40 cm) above patterns of different spatial wavelength (1.8–90°) in the direction of flight. The OFM modulate their flight speed three‐fold over different patterns. They fly fastest when there is no pattern in the tunnel or the contrast elements are too narrow to resolve. When the spatial wavelength of the pattern is sufficiently wide to resolve, moths fly at a speed that tends to maintain a visual contrast frequency of 3.5 ± 3.2 Hz rather than a constant angular velocity, which varies from 57 to 611° s^?1. In addition, for the first time, it is also demonstrated that the width of a contrast pattern perpendicular to the flight direction modulates flight speed.     

Adjustment of flight speed of gregarious desert locusts (Orthoptera: Acrididae) flying side by side

In tethered flying locusts, optomotor thrust responses induced by translatory pattern motion within the lateral visual fields were studied under closed-loop conditions. By modulating thrust in a compensatory manner, locusts counteracted a bias motion superposed on the thrust-related motion. This way, pattern speed was kept at 0° s^–1, indicating the set point of the respective optomotor control circuit. Though the quality of bias compensation varied greatly, it was largely independent from pattern characteristics. It might indicate that the gain of behavior not only is controlled by an automatic mechanism but also is affected by spontaneous modulations. Compensation of bias motion was critically dependent on the relation between self- and bias-generated motion: Locusts did not take control over pattern motion if self- and bias-generated motion differed greatly. Instead, locusts adopted a constant, supposingly preferred, thrust value. Therefore, flight speed is assumed to be controlled by two systems: the optomotor and a preferred thrust system. In free flight, an equalization of the flight speed of locusts within a swarm might result from similar behavior. In combination with a presumed coordination of the locusts' course direction, this may explain the continued cohesion of swarms in the field.     

Optomotor regulation of ground velocity in moths during flight to sex pheromone at different heights

ABSTRACT. Males of two species of moths ( Grapholitha molesta (Busck) and Heliothis virescens (F.)) were flown in a sustained-flight tunnel in horizontal pheromone plumes. The up-tunnel velocity of the moths increased with increasing height of flight and for G.molesta was independent of tunnel wind velocities. Use of moving ground patterns verified that the height of flight above the ground was the factor related to the changes in up-tunnel velocity. Even though up-tunnel velocity increased with increased flight height, angular velocity of image motion did not. Males appeared to use visual cues from the ground pattern and from other sources to determine their up-tunnel velocities. The relationship of preferred retinal velocities to optomotor anemotaxis is discussed.     

Control of flight by means of lateral visual stimuli in gregarious desert locusts, Schistocerca gregaria

Abstract. Tethered flying locusts were stimulated either by a periodic grating or by a spotted 'swarm-simulating' pattern moving horizontally, parallel to their longitudinal body axis within their lateral visual fields. The direction of movement of the pattern was changed periodically from progressive to regressive and vice versa.
Both kinds of patterns induced a correlated modulation of yaw-torque and thrust. The two measured flight parameters were modulated independently of each other. Each parameter either increased with progressive and decreased with regressive pattern motion or vice versa. The characteristic curves of thrust and yaw-torque responses - i.e. response amplitude versus contrast frequency resp. angular velocity – measured upon stimulation with the periodic grating between 2 and 70 Hz were at a maximum at 10 Hz and decreased at higher and lower contrast frequencies. The shape of the curves was nearly identical. The characteristic curves measured upon stimulation with the 'swarm-simulating' pattern between 60 and 1500^o s^-1 could be simulated using the spatial wavelength content of the pattern and the characteristic curves for periodic gratings.
Therefore, we suggest that the speed and direction of locusts' flight result from the optomotor effectiveness of the pattern image formed by the neighbouring individuals under free flight. The measured responses would thus contribute to the common orientation of groups of locusts within a migrating swarm and thus to swarm cohesion.     

The kinematics of Heteroptera in free flight

The kinematics of six species of Heteroptera in free flight are analysed and compared.

• (1) 

  Using nested analysis of variance techniques, statistically significant variation was detected between species for several of the flight parameters measured: mean angular velocity; pronation/supination ratio; upstroke/downstroke ratio; and wing beat frequency. In each case this is discussed in terms of variation in flight behaviour.
• (2) 

  Beneficial aerodynamic forces are generated during the upstroke and the downstroke, in both fast forward and rising flight.
• (3). 

  When the insects change from level, forward flight to near vertical, rising flight, the following parameters are altered in most of the sequences analysed:

• (a). 

  the stroke plane angle becomes steeply, negatively inclined, associated with an increase in body angle;
• (b). 

  the stroke amplitude is reduced;
• (c). 

  wing beat frequency is lowered, associated with a drop in mean angular velocity;
• (d). 

  the speed of stroke reversal (rotational velocity) is increased. This may be associated with increased wing torsion and tip flexion which in turn could improve any beneficial unsteady aerodynamic effects generated at stroke reversal.

The reasons for this change in flight performance and the deviations from that seen in other insects are discussed.
It is shown that Heteroptera may make use of wing drag in flight, particularly during rising flight.     

Flying Slower: Floor Pattern Object Size Affects Orthokinetic Responses During Moth Flight to Sex Pheromone

Previous studies with Oriental Fruit Moth (OFM, Grapholita molesta) and Heliothis virescens males flying upwind along a pheromone plume showed that they increased their upwind flight speed as they flew higher above striped floor patterns and, for OFM, to a similar degree over dotted floor patterns. This response pattern has been demonstrated in another moth species, Epiphyas postvittana and in a beetle, Prostephanus truncatus. In all cases the role played by the change in angular size of the wind tunnel’s ventral floor pattern was not assessed. In the present study we specifically addressed this question with a systematic examination of moths’ flight control over different sizes of transverse stripes and dot patterns ranging down by halves from 5 to 0.625 cm and a blank white floor as a control, and showed that OFM males fly faster upwind and along their flight paths over floor patterns of decreasing size. Increased speeds over striped patterns were evident as stripe width decreased below 2.5 cm, whereas moths did not increase their flight speed over dot patterns until dot size had decreased to less than 1.25 cm. Another flight component that the moths can actively control, their course angles, was unchanged above both patterns, except for moths flying over 5 cm stripes. Turning frequency and interturn distances were mostly unchanged or offset each other, negating any effects on upwind progress. As in an earlier study examining flight speeds at three heights above floor patterns of three densities, the moths’ changes in speed appear to be exclusively affected by changes in their orthokinetic response to the size of the floor pattern objects.     

Basic organization of operant behavior as revealed in Drosophila flight orientation

Summary Operant behavior is studied in tethered Drosophila flies using visual motion, heat or odour as operandum and yaw torque, thrust or direction of flight as operans in various combinations (Fig. 1). On the basis of these results a conceptual framework of operant behavior is proposed: (1) It requires a goal (desired state) of which the actual state deviates. (2) To attain the goal a range of motor programs is activated (initiating activity, see Fig. 7). (3) Efference copies of the motor programs are compared to the sensory input referring to the deviation from the desired state (e.g. by cross-correlation). (4) In case of a significant coincidence the respective motor program is used to modify the sensory input in the direction towards the goal. (5) Consistent control of a sensory stimulus by a behavior may lead to a more permanent behavioral change (conditioning). In this scheme operant activity (1–4) and operant conditioning (1–5) are distinguished.Abbreviations ALU arbitrary length unit - d horizontal angular width of visual pattern - IR infrared - SEM standard error of the means - T yaw torque - Th thrust - performance index - horizontal angle between visual pattern position and longitudinal body axis of the fly - vertical angular extension of visual pattern     

Elementary pattern discrimination (behavioural experiments with the fly Musca domestica)

This paper investigates the problem of spontaneous pattern discrimination by the visual system of the fly. The indicator for discrimination and attractivity of a pattern is the yaw torque of a test fly. It is shown that the pattern discrimination process may be treated as a special (degenerate) case of figureground discrimination which has been described in detail in earlier publications. Decisive for the discrimination process is the fact that pattern discrimination by the fly is mediated by motion detectors which respond not only a pattern velocity but also to structural properties of pattern contrast. This is demonstrated by the transition from the existing twodimensional array of motion detectors to a continuous detector field which enabled us to calculate instantaneous detector responses to instationary pattern motion. The new approach, together with the special theory for figure-ground discrimination, is then applied to predict spontaneous discriminations of onedimensional periodic patterns. It is shown that predictions and experimental results are in good agreement. The second set of discrimination experiments deals with two dimensional dot patterns for which a quantitative theory is not yet available. However, it is shown that the attractivity of a dot pattern crucially depends on both the orientation and the direction of motion relative to the fly's eyes. If the contrast of a moving dot elicits an event in a motion detector which through the detector's time constant leads to an interference with an event received by a preceeding dot, the attractivity of the dot pattern is diminished. In the discussion relations are drawn between the concepts of pattern discrimination in honey bees and the theoretical aspects of discrimination put forward in this paper. It is briefly discussed why a two-dimensional motion detector theory might become the key for an understanding of pattern categories like figural intensity and figural quality.     

Visual control of straight flight in Drosophila melanogaster

Summary The optomotor system of Drosophila is investigated in a flight simulator in which the fly's yaw torque controls the angular velocity of the panorama (striped drum, negative feedback). Flies in the flight simulator maintain a stable orientation even in a homogeneously textured panorama without landmarks. During straight flight, torque is not zero. It consists of small pulses mostly alternating in polarity. The course is controlled by the duration (and possibly amplitude) of the pulses. The system operates under reafference control. By comparing the pulses with the visual input the system continuously measures and adjusts the efficacy of the torque output. The comparison, however, is not between angular velocity and yaw torque but, instead, between visual acceleration and pretorque, the first time derivative of torque. For comparison, the system first computes a cross-correlation. If the correlation coefficient is above a certain threshold the system calculates the external gain and adjusts its internal gain so as to keep the total gain constant. With the correlation coefficient below threshold, however, the system keeps the internal gain low despite the infinitely small external gain. We propose that for a reafferent optomotor system the coupling coefficient and the correlation coefficient of pretorque and visual acceleration are more relevant than the distinction between exafference and reafference.Abbreviation EMD elementary movement detector     

Directional control by male gypsy moths of upwind flight along a pheromone plume in three wind speeds

By attaching a reflective strip to the thorax, we documented with video recordings in a wind tunnel the spatial orientation of male gypsy moths, Lymantria dispar, as they flew along a plume of sex pheromone. In wind speeds of 61, 122, and 183 cm s⁻¹, moths flew very similar tracks along a pheromone plume. Moths aimed their thrust closer to upwind in increasing wind speeds using a roll maneuver. As a result, the orientation of their visual flow field, represented by the slip angle (the angular distance between the direction of flight and the longitudinal body axis), remained relatively constant. We propose that directional control during self-steered zigzagging is achieved by rolling, thereby maintaining a set slip angle. Following a roll at the apex of a turn that aligns the moth with its preferred slip angle, a moth banks toward a cross wind leg. By banking moths may maintain a stable image flow at oblique angles to their longitudinal body axis. Accepted: 16 July 1998     

Chasing and pursuit in the dolichopodid fly Poecilobothrus nobilitatus

1. Male Poecilobothrus nobilitatus show two distinct kinds of pursuit. Females are “shadowed” at a distance of a few cm, using both rotational and lateral movements. Other males are chased, in a pursuit that involves only rotation and fast forward flight. The rotational component of pursuit appears to have the same control system in both types of tracking, and it is best described as a continuous translation of the error angle between the direction of the target and the pursuing fly's body axis into the pursuing fly's angular velocity. The constant of proportionality is 30–40°·s^?1 per degree, and the delay in the system is about 15 ms. Pursuit on the ground is 2–3 times slower than in flight, although the delay seems to be similar.
2. Attempts were made to see whether the aerial pursuits could be modelled effectively by a saccadic or discontinuous control system, as suggested for Musca pursuit (Wagner 1986). It was found that the velocity profiles of the chases could be fitted by an overlapping series of plausible saccade-like events, However, the correlation between visual information (error angle and error angular velocity) available just before each fictive saccade correlated poorly with saccade peak velocity. It is thus concluded that Poedlobothrus pursuit is basically continuous in nature, but it is argued that the two types of mechanism are hard to distinguish in natural behaviour.

     

Optomotor control of wing beat and body posture in drosophila

Continuous movement of striped patterns was presented on either side of a tethered fruitfly, Drosophila melanogaster, in order to simulate the displacement of stationary landmarks within the visual field of the freely moving fly. The horizontal components of the stimulus elicit, predominantly, yaw-torque responses during flight, or turning responses on the ground, which counteract involuntary deviations from a streight course in the corresponding mode of locomotion. The vertical components elicit, predominantly, covariant responses of lift and thrust which enable the fly to maintain a given level of flight. Monocular stimulation is sufficient to produce antagonistic responses, if the direction of the stimulus is reversed. The following constituents of the responses were derived mainly from properties of wing beat and body posture on photographs of fixed flight under visual stimulation. Wing stroke modulation (W. S. M.): The difference, and the sum, of the stroke amplitudes on either side are independently controlled by horizontal and vertical movement components, respectively. The maximum range of modulation per wing (12.3°) is equivalent to a 63% change in thrust on the corresponding side. Leg stroke modulation (L.S.M.): In the walking fly each pair of legs is under control of visual stimulation. The details of leg articulation are still unknown. Abdominal deflection (A.D.): An actively induced posture effect. Facilitates steering during free flight at increased air speed. Hind leg deflection (H.L.D.): Same as before. On most of the photographs the hind legs were deflected simultaneously and in the same direction as the abdomen. Hitch inhibition (H.I.): The term hitch denotes a transient reduction of stroke amplitude which seems to occur spontaneously and independently on either side of the fly. The hitch angle (12.2±3.8° S.D.) is most probably invariant to visual stimulation. Hitches are comparatively frequent in the absence of pattern movement. Their inhibition under visual stimulation is equivalent to an increase of the average thrust of the corresponding wing. The different constituents contribute to the optomotor responses according to the following tentative scheme (Fig. 7). The torque response is essentially due to the effects of W.S.M., A.D., H.L.D. and H.I., and the turning response to L.S.M. and possibly H.L.D., if the landmarks drift from anterior to posterior. So far, H.I. seems to be the only source of the torque response, and L.S.M. the only source of the turning response, if the landmarks drift in the opposite direction. The lift/thrust response results essentially from the effects of W.S.M. and H.I., no matter whether the landmarks drift from inferior to superior or in the opposite direction. The results obtained so far suggest that the optomotor control of course and altitude in Drosophila requires at least eight independent input channels or equivalent means for the separation of the descending signals from the visual centres. Further extension and refinement of the wiring scheme is required in order to improve the identification of the sensory inputs of the motor system and the classification of optomotor defective mutants.     

Monitoring watercourse vegetation, a synecological approach to dynamic gradients

Changes in vegetation under reduced control measures over 3 to 5 years in watercourses in a rural environment in The Netherlands were evaluated. A method to deal with slow changes on a steep gradient is presented. The gradient with various vegetation types between the middle of the watercourse and the bank-top was split up into zones. Species composition of each zone was evaluated using literature on syntaxonomy. Cover of character species, multiplied by the width of the zones, was used to quantify the contribution of various syntaxa in the vegetation. Changes in these contribution data were used to evaluate changes over the years. The method was applied to two experiments in which cleaning frequency was reduced. Submerged vegetation of Callitriche-Ranunculetum penicillati in one and of Potamogetonetalia pectinati in the other case, hardly changed. Emergent vegetation of Nasturtio-Glycerietalia or Sparganio-Glycerietum fluitans tended to expand into the submerged zone. Bank vegetation began to show signs of development into ruderal vegetation, as a shift from Molinio-Arrhenatheretea into Artemisietea was detected. The method allowed the conclusion that conditions were too eutrophic in both experiments for a diverse brook vegetation development without additional habitat improvement.     

风洞内粘虫飞翔行为与气流的关系

利用自制的昆虫飞翔实验风洞 ,系统观测了在风洞条件下粘虫在不同流速实验气流中的起飞行为与飞翔行为。结果表明 ,微风能刺激粘虫起飞 ,试虫表现明显的偏爱迎风 (或稍偏一点角度 )起飞的习性 ,飞翔时亦多采取迎风 (或稍偏一点角度 )的姿势。试虫在气流中的实际位移是昆虫飞翔位移与气流位移的矢量和。当气流速度小于 2 m/ s时 ,逆风向位移占多数 ;而气流速度为 3～ 4 m/ s时 ,94 .8%的试虫为顺风向位移。     

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.     

Variability in odor-modulated flight by moths

Based on previous studies of odor-modulated flight where track parameter data was lumped and averaged, the speed and orientation of the moths' movement along their flight tracks have been said to be controlled to maintain certain “preferred” values. The results from our fine-scaled analysis of this behavior show that none of the track parameters typically measured are held constant. The moths' speed along the flight track is modulated substantially and predictably: fastest along the straight legs and slowest around the turns. In addition, about half of the individuals studied progressively reduced the peak speed along the straight legs as they approached the pheromone source. While most of the track legs between the turns were directed upwind, their orientations were widely distributed, indicating no preferred direction. Small fluctuations of orientation along some straight legs suggest corrective maneuvers to stabilize flight direction about an internal set point. The visual inputs hypothesized to control steering and speed, transverse and longitudinal image flow, changed continuously during upwind flight in pheromone, but no regular relationship between them was observed. We found that the orientation of the longitudinal body axis and the direction of thrust (course angle) were only rarely coincident during upwind flight to the odor source, suggesting that moths receive sensory input which differs quantitatively from that calculated by conventional methods. Our results strongly suggest that the long-accepted hypothetical mechanisms of control for this behavior do not operate in the manner in which they have been proposed. Accepted: 11 July 1997     

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.     

Optic flow-based collision-free strategies: From insects to robots

Flying insects are able to fly smartly in an unpredictable environment. It has been found that flying insects have smart neurons inside their tiny brains that are sensitive to visual motion also called optic flow. Consequently, flying insects rely mainly on visual motion during their flight maneuvers such as: takeoff or landing, terrain following, tunnel crossing, lateral and frontal obstacle avoidance, and adjusting flight speed in a cluttered environment. Optic flow can be defined as the vector field of the apparent motion of objects, surfaces, and edges in a visual scene generated by the relative motion between an observer (an eye or a camera) and the scene. Translational optic flow is particularly interesting for short-range navigation because it depends on the ratio between (i) the relative linear speed of the visual scene with respect to the observer and (ii) the distance of the observer from obstacles in the surrounding environment without any direct measurement of either speed or distance. In flying insects, roll stabilization reflex and yaw saccades attenuate any rotation at the eye level in roll and yaw respectively (i.e. to cancel any rotational optic flow) in order to ensure pure translational optic flow between two successive saccades. Our survey focuses on feedback-loops which use the translational optic flow that insects employ for collision-free navigation. Optic flow is likely, over the next decade to be one of the most important visual cues that can explain flying insects' behaviors for short-range navigation maneuvers in complex tunnels. Conversely, the biorobotic approach can therefore help to develop innovative flight control systems for flying robots with the aim of mimicking flying insects’ abilities and better understanding their flight.     

A model of visually-guided smooth pursuit eye movements based on behavioral observations

We report a model that reproduces many of the behavioral properties of smooth pursuit eye movements. The model is a negative-feedback system that uses three parallel visual motion pathways to drive pursuit. The three visual pathways process image motion, defined as target motion with respect to the moving eye, and provide signals related to image velocity, image acceleration, and a transient that occurs at the onset of target motion. The three visual motion signals are summed and integrated to produce the eye velocity output of the model. The model reproduces the average eye velocity evoked by steps of target velocity in monkeys and humans and accounts for the variation among individual responses and subjects. When its motor pathways are expanded to include positive feedback of eye velocity and a switch, the model reproduces the exponential decay in eye velocity observed when a moving target stops. Manipulation of this expanded model can mimic the effects of stimulation and lesions in the arcuate pursuit area, the middle temporal visual area (MT), and the medial superior temporal visual area (MST).     

Visual edge orientation shapes free-flight behavior in Drosophila

Insects rely on visual cues to estimate and control their distance to approaching objects and their flight speed. Here we show that in free-flight, the motion cues generated by high-contrast vertical edges are crucial for these estimates. Within a visual environment dominated by high-contrast horizontal edges, flies fly unusually fast and barely avoid colliding with the walls of the enclosure. The disruption of flight behavior by horizontal edges provides insight into the structure of visually-mediated control algorithms.