Height control by free-flying Drosophila

ABSTRACT. In a horizontal wind tunnel, Drosophila flew at almost constant height along tracks up to 2 m long. The flies rose or sank only slowly when it was so dark that they no longer responded to movements of the tunnel floor, suggesting that their height control is mediated, at least partly, by responses to their movement relative to the air. In the light, the flies maintained height better than in the dark and were very responsive to movements around them. They faithfully followed the up and down movements of horizon screens at their sides whether they were flying in still air or against a wind, even in the presence of many other stationary visual cues. The flies did not respond by compensatory height changes to real vertical movements of a patterned horizontal disc beneath them, nor to changes in the size of the floor pattern. They did respond to horizontal acceleration of the floor pattern in the direction opposite to their flight (optically simulating a descent by the fly), by an apparently compensatory increase in height, but they also rose (instead of sinking) in response to floor acceleration in the direction of their flight. When the floor was accelerated in either direction they showed compensatory groundspeed-controlling responses. The increases in height might be alarm responses to sudden movements in the visual field beneath them. Both speed and height changing responses to floor movement were reduced when the number of stationary visual cues was increased. Drosophila thus control their height mainly by responses to the apparent movement of nearby visual cues at round about their own height.     

On the encoding of movement vectors by honeybees. Are distance and direction represented independently?

Honeybees flying repeatedly over the same trajectory link it to an associated visual stimulus such that on viewing the stimulus they perform a trajectory in the habitual direction. To test if trajectory length can also be linked to a visual stimulus, bees were trained to fly through a multi-comparmented maze. Bees flew through a multi-compartmented maze. In one compartment a short trajectory could be linked to a stripe pattern oriented at 45° to the horizontal. In another compartment a longer trajectory could be linked to 135° stripes. Bees made both associations: their trajectories were short when viewing 45° stripes and longer when viewing 135° stripes. 90° stripes evoked trajectories of intermediate length.To test if distance and direction are linked independently to stripe orientation, a bee's trajectory was linked to 135° stripes in one compartment and to 45° stripes in another. These trajectories were the same length but differed in their horizontal direction by 60° or by 120°. 90° stripes evoked trajectories of intermediate direction which were shorter than those elicited by either training pattern. Bees were also trained to generate one long and one short trajectory with directions 120° apart. The trajectories elicited by 90° stripes were then biased towards the direction of the long training vector. Length and direction are not treated separately. The rules for combining trajectories resemble those of vector averaging.     

Do tsetse flies ‘see’ zebras? A field study of the visual response of tsetse to striped targets

Abstract A field study in Zimbabwe of Glossina pallidipes Austen and G. morsitans morsitans Westwood supported Waage's (1981) hypothesis that the striped pattern of zebras may protect them from being bitten by blood-sucking flies. In addition, the results suggest that the orientation of the stripes may be crucially important for the unattractiveness of zebras. The relative attractiveness of five different stationary targets (black, white, grey, vertically-striped and horizontally-striped; stripe width = 5 cm) were each tested on their own and in pairs of all combinations, with artificial host odour (CO₂ plus acetone) always present. Electric nets were used to catch flies as they attempted to land on or circle the targets. The results were similar for the two species of tsetse. When tested on their own, grey and vertically-striped targets caught similar numbers of flies and both caught significantly fewer than black or white targets (c. 36% as many). Horizontally-striped targets caught <10% as many flies as any other single target. Although there was no significant difference between the attractiveness of grey and vertically striped targets when they were presented together, when paired with the other targets, grey was as attractive as black or white, but the vertically-striped target was significantly less attractive than black or white (P < 0.001). In other words, a difference between grey and vertical stripes was found only in their attractiveness in relation to other targets. The horizontally-striped target, however, always caught the fewest flies, regardless of whether it was presented alone or alongside another target.     

The generalization of directional visual stimuli in the honey bee, Apis mellifera

In transfer tests the ability of bees to generalize visual stimuli was tested by using differently inclined stripes and stripe patterns offered on a vertical screen. After having been trained to single stripes or equidistant stripe patterns, which were orientated by α₊ = 45° to the horizontal, the bees had to discriminate between the training direction α₊ and the competition direction α_c = 135° by means of special stripe configurations. These transfer patterns were obtained by varying different stimulus parameters of the original training stripes, for example by (1) reversing contrast between a stripe and the surrounding visual field, (2) changing the ratio of length/width and by this the dimensions of the stripe, and (3) inserting white intervals into the black stripes. In all three test series the bees succeeded in detecting the α₊-direction along a broad range of stimulus variations. As the bees in the transfer tests positively responded to patterns, which on the other side were significantly discriminated from the training pattern (control tests), the information about the direction of the visual cue had been transferred to a new pattern configuration never seen by the bees during the training situation.     

A behavioural test of the sensitivity of a nocturnal mosquito, Anopheles gambiae, to dim white, red and infra-red light

Abstract. A behavioural test was used to determine the light sensitivity of the nocturnal mosquito Anopheles gambiae Giles s.s. to low intensities of 'white' light (tungsten filament), 'red' light (white light filtered by a darkroom safelight filter) and 'infra-red' light) of two types (white light filtered by a λ>700 nm filter, and light-emitting diodes with λ>900 nm). Mosquitoes were placed in a 20 cm diameter flight-tunnel and their 'optomotor' response to a pattern of stripes moving across their visual field (at 14.5 cm s^-1) was recorded with infra-red-sensitive video. In free-flight, with ample light, the mosquitoes controlled their flight speed and direction in relation to the stripe movement, so that the stripes always appeared to move across their visual field from front to back. They did this by flying either with the moving stripes fast enough to overtake them (19.5 ± 0.7 cm s^-1), or against them more slowly (10.3 ± 0.7 cm s^-1)- The net ground speed of the mosquitoes was thus c. 4–5 cm s^-1. This response was significant down to 10^-5 W m^-2 in 'white' light, and 10^-3 W m^-2 in 'red' light. At light intensities below threshold and in infra-red light, however, they appeared to fly at random with respect to the stripe movement. The assumption commonly made, that mosquitoes do not 'see' in red light, may thus have to be revised.     

Fixation and optomotor response of walking colorado beetles: interaction with spontaneous turning tendencies

Abstract The paths of Colorado beetles ( Leptinotarsa decemlineata Say) in a featureless environment are circular, like those of other species studied. The turning velocity may reach 35^o/s and is due to an internal asymmetry, which may change spontaneously. Normally, all control loops of the insect, like fixation or optomotor responses, must work against this asymmetry to stabilize the insect's path. Stationary vertical patterns damp this turning tendency, but their effect is not strong enough to induce a straight path. Only 70% of the turning tendency can be so eliminated. This reaction is termed optomotor response because it can be adequately described with the parameter turning velocity alone. The insect's path was stabilized more effectively when pattern wavelengths were greater than 60^o. The insects seemed to fixate these wider stripes. This reaction is termed fixation because the correlation between pattern components and insect's course becomes prominent.
A comparison was made between these reactions to stationary patterns and to turning patterns. No differences could be found in the behavioural reactions to the different situations. This suggests that the insect does not use an internal representation of its spontaneous turning tendency to discriminate between the type of turning of the optical environment. These results can be explained by a simple feedback control loop with an additive interaction between the internal turning command and feedback signals from the eyes.     

Movement learning of free flying honeybees

Summary Free flying honeybees were conditioned to moving black and white stripe patterns. Bees learn rapidly to distinguish the direction of movement in the vertical and horizontal plane.After being trained to a moving pattern bees do not discriminate the moving alternative from a stationary one. There is no significant velocity discrimination for patterns moving in the same direction.For vertical movements there are clear asymmetries in the spontaneous choice preference and in the learning curves for patterns moving upward or downward.After bees are trained to a stationary pattern they can discriminate it from an upward moving alternative. Learning curves involving movement are generally biphasic, suggesting different adaptive systems depending on the number of rewards.The flight pattern of bees which are trained to movement changes during the process of learning. At the beginning of the learning procedure bees reveal an optokinetic response to the moving patterns, this response is strongly reduced after a number of rewards on a moving pattern.     

Non-periodic cues generate seven ftz stripes in the Drosophila embryo

We have examined the expression pattern of the segmentation gene fushi tarazu (ftz) by in situ hybridization to whole mount embryos using digoxygenin labeled probes. This method has revealed previously undetected stages in the development of the ftz RNA pattern. The ftz stripes arise individually in a distinct, non-linear order along the anterior-posterior axis of the embryo. In addition, the stripes develop differentially along the dorsal-ventral axis; most stripes emerge on the ventral side and then gradually spread dorsally until they surround the entire circumference of the embryo. The order of appearance of ftz stripes is not inversely correlated with the order of appearance of hairy (h) stripes as would be expected if ftz stripes were generated by h repression. Furthermore, the seven ftz stripes are correctly established in embryos carrying mutations in h, eve or runt, with normal expression patterns decaying only after cellularization. Thus, the so called primary pair-rule genes are involved in the refinement rather than establishment of the ftz stripes. The contribution of cis-acting regulatory elements to the ftz pattern was examined. The zebra and upstream elements interact to generate seven correctly positioned stripes at the end of cellularization. However, stripe establishement is not correctly mimicked by any ftz/lac fusion gene: stripes arise in an order drastically different from the endogenous ftz gene suggesting the existence of ftz regulatory elements outside the 10-kb region examined to date. These observations suggest that the ftz pattern is directed by at least two independent regulatory systems: first, stripe establishment is directed by regionally distributed factors that act differentially in individual stripes along both anterior-posterior and dorsal-ventral axes of the egg and, second, stripe refinement and maintenance are mediated by pair-rule gene products that interact with previously identified ftz regulatory elements. This multi-level regulation provides a back-up system that ensures the development of seven stripes in the blastoderm.     

Spatial distribution of Prosopis glandulosa var. torreyana in vegetation stripes of the southern Chihuahuan Desert

Mosaics consisting of vegetation stripes surrounded by bare areas have been described in several arid and semiarid ecosystems. The dynamics of the system depends on the redistribution of rainwater which is preferentially stored and evapotranspired in the vegetated stripes. A process of plant `colonization' in the upslope fringe of the stripes has been described in some cases and a consequent upslope migration of the stripes has been inferred, but not confirmed in all cases quoted in the literature. In this paper, we studied the spatial distribution of mesquite (Prosopis glandulosa var. torreyana) and the soil parameters in three vegetation stripes and their associated bare areas in the southern Chihuahuan Desert. The spatial distribution of mesquites of different sizes do not coincide with that expected under the hypothesis of an uniform upslope stripe migration, but soil data suggest that current bare areas had been vegetated some time ago. Dispersion and establishment abilities enhanced by overgrazing may explain the observed mesquite distribution, but the presence of trees with high basal diameters in any part of the stripes suggests stripe permanence at the same site and no upslope migration. These results point to the conflicting evidence on stripe migration that has been already found in other areas. The most probable scenario in our study area is that of a general long-term change of form of the stripes taking place at very variable speeds in different stripes, including the possibility that some of them remain stationary for prolonged periods, and showing different histories of colonization according to the life-history of the different species concerned. The speed and regularity of the process would show a very high temporal and spatial variability due to the interaction of climatic, geomorphologic and biotic interactions.     

Pattern formation during insect leg segmentation: Studies with a prepattern of a cell surface antigen

Summary A monoclonal antibody (MAb) that binds to a cell surface antigen selectively localized to epithelial cells undergoing morphogenesis was used to study the segmentation of the growing embryonic leg of the cockroachPeriplaneta americana. The MAb labels circumferential stripes of cells at locations where invagination will occur to form the leg segments. The formation of these stripes precedes any morphological change in the epithelial layer or in individual cells. The temporal and spatial distribution of the antigen indicates the existence of a prepattern for leg segmentation, examination of which can give information about pattern generating mechanisms. Although highly stereotyped, the sequence in which the stripes appear does not follow a simple pattern proceeding in one direction along the proximal-distal axis. It is proposed that each stripe is a boundary in a positional field. Stripe formation leads to the division of the leg into a repeating series of identical positional fields. Three different mechanisms for the formation of stripes of MAb labeled cells have been observed and the role of each in the evolution of the insect leg is discussed. Measurements of leg and leg segment lengths when the various stripes appear has demonstrated considerable variation, particularly at the early stages of segmentation. Rules or mechanisms generating pattern at early stages of development are not rigid. Variations arising are compensated for by later occurring events so that stereotyped structures are formed.     

Size-dependent pigmentation-pattern formation in embryos of Alligator mississippiensis: time of initiation of pattern generation mechanism

The pigmentation pattern of Alligator mississippiensis was examined. The number of white stripes on the dorsal side of embryos (stages 21-28) and hatchlings from eggs incubated at 30 degrees C (100% females) and 33 degrees C (100% males) was recorded. Total length, nape-rump length and tail length were recorded for each embryo and hatchling. The number of white stripes was affected by incubation temperature but not sex; hatchlings incubated at 33 degrees C had two more white stripes than those at 30 degrees C, despite being the same length. Five female hatchlings produced at 33 degrees C by manipulation of the temperature, had the same number of stripes as males that developed under the same incubation temperatures. The appearance of the pigmentation was accelerated in embryos incubated at 33 degrees C, occurring eight days earlier than at 30 degrees C. At the time just before the first signs of pigment deposition, embryos from 33 degrees C were longer than those at 30 degrees C. If the stripe formation is size dependent this explains why hatchlings at 33 degrees C have more stripes than hatchlings from 30 degrees C. The mechanism that produces the stripe patterns is unknown. We describe key elements a pattern formation mechanism must possess to produce such stripes and suggest a possible mechanism, based on cell movement driven by chemotaxis. We apply the mathematical model to dorsal patterning on A. mississippiensis. We show how length at pattern formation is the prime factor in determining stripe number and how the pattern can be formed in the observed anterior-posterior sequence. We present numerical simulations and show that the qualitative behaviour is consistent with the experimental results.     

Head movements of the mealworm beetle Tenebrio molitor

Tethered walking imagines of the mealworm beetle Tenebrio molitor wave their heads in random fashion. If a periodic pattern of vertical black and white stripes is rotated around the animal a regular nystagmic head movement is superimposed upon the random waving, the frequency of the latter equals the contrast frequency within large ranges of the angular velocity of the pattern. The nystagmus is inverted: After a short period of tracking, during which the angular velocity of the head is the same as that of the panorama, the head returns slowly toward its normal position according to an exponential-like function. Resting animals do not wave their heads. However, if the above panorama is rotated, the beetle turns its head in the direction of the movement of the panorama and holds it in a side-way position, as long as the rotation is maintained. The angular position reached depends in the same manner on the angular velocity of the panorama as the turning tendency of walking animals established in open loop experiments using the spherical Y-maze method.     

Mechanisms of black and white stripe pattern formation in the cuticles of insect larvae

Molecular mechanisms that produce pigment patterns in the insect cuticle were studied. Larvae of the armyworm Pseudaletia separata have stripe patterns that run longitudinally along the body axis. The pattern in the cuticle became clear by being emphasized by the increasing contrast between the black and white colors of the lines after the last larval molt. We demonstrated that dopa decarboxylase (DDC) mRNA as well as protein are expressed specifically in the epidermal cells under the black stripes. The pigmentation on the stripes was clearly diminished by injection of a DDC inhibitor (m-hydroxybenzylhydrazine) to penultimate instar larvae for 1 day before molting, suggesting that DDC contributes to the production of melanin. Further, electron microscopic observation showed that the epidermal cells under the gap cuticle region (white stripe) between the black stripes contain many uric acid granules, which gives a white color. Our findings suggest that the spatially regulated expression of DDC in the epidermal cells produces the black stripes while abundant granules of uric acid in the cells generate the white stripes in the cuticle. Based on these results, we concluded that this heterogeneity in the epidermal cells forms cuticular stripe patterns in the armyworm larvae.     

Habitat corridors function as both drift fences and movement conduits for dispersing flies

Corridors connect otherwise isolated habitat patches and can direct movement of animals among such patches. In eight experimental landscapes, we tested two hypotheses of how corridors might affect dispersal behavior. The Traditional Corridor hypothesis posits that animals preferentially leave patches via corridors, following them into adjacent patches. The Drift Fence hypothesis posits that animals dispersing through matrix habitat are diverted into patches with corridors because they follow corridors when encountered. House flies (Musca domestica L.), a species that prefers the habitat of our patches and corridors, were released in a central patch (100×100 m) and recaptured in peripheral patches that were or were not connected by a corridor. Flies were captured more frequently in connected than unconnected patches, thereby supporting the Traditional Corridor hypothesis. The Drift Fence hypothesis was also supported, as flies were captured more frequently in unconnected patches with blind (dead end) corridors than in unconnected patches of equal area without blind corridors. A second experiment tested whether these results might be dependent on the type of patch-matrix boundary encountered by dispersing flies and whether edge-following behavior might be the mechanism underlying the observed corridor effect in the first experiment. We recorded dispersal patterns of flies released along forest edges with dense undergrowth in the forest (“closed” edges) and along edges with little forest understory (“open” edges). Flies were less likely to cross and more likely to follow closed edges than open edges, indicating that when patch and corridor edges are pronounced, edge-following behavior of flies may direct them along corridors into connected patches. Because edges in the first experiment were open, these results also suggest that corridor effects for flies in that experiment would have been even stronger if the edges around the source patches and corridors had been more closed. Taken together, our results suggest that corridors can affect dispersal of organisms in unappreciated ways (i.e., as drift fences) and that edge type can alter dispersal behavior.     

Fast and slow flight torque responses in flies and their possible role in visual orientation behaviour

The flight torque responses of tethered flying houseflies to motion and presentation or removal of a vertical dark stripe on a bright background were recorded in real time. Motion with constant speed of 100° s^-1 from front to back elicits a strong fast response following the diraction of the stimulus motion. Motion from back to front elicits a weaker response. Instantaneous presentation and removal of a stationary stripe elicit weak, slow response. Apparent motion from front to back and from back to front elicit weak responses with a fast, directionally selective, transient peak followed by a slow response component oriented towards the stripes position. The fast transient peak response is not elicited if the animals were stimulated before with real movement of the stripe. The results are discussed and an earlier proposed model for free flight tracking and fixation is extended.     

Correlation between the ripple phase and stripe domains in membranes

We investigate the relationship between stripe domains and the ripple phase in membranes. These have previously been observed separately without being linked explicitly. Past results have demonstrated that solid and ripple phases exhibit rich textural patterns related to the orientational order of tilted lipids and the orientation of ripple corrugations. Here we reveal a highly complex network pattern of ripple and solid domains in DLPC, DPPC bilayers with structures covering length scales from 10 nm to 100 μm. Using spincoated double supported membranes we investigate domains by correlated AFM and fluorescence microscopy. Cooling experiments demonstrate the mode of nucleation and growth of stripe domains enriched in the fluorescent probe. Concurrent AFM imaging reveals that these stripe domains have a one-to-one correspondence with a rippled morphology running parallel to the stripe direction. Both thin and thick stripe domains are observed having ripple periods of 13.5±0.2 nm and 27.4±0.6 nm respectively. These are equivalent to previously observed asymmetric/equilibrium and symmetric/metastable ripple phases, respectively. Thin stripes grow from small solid domains and grow predominantly in length with a speed of ~3 times that of the thick stripes. Thick stripes grow by templating on the sides of thinner stripes or can emerge directly from the fluid phase. Bending and branching angles of stripes are in accordance with an underlying six fold lattice. We discuss mechanisms for the nucleation and growth of ripples and discuss a generic phase diagram that may partly rationalize the coexistence of metastable and stable phases.     

Similarities and differences in the peering-jump behavior of three grasshopper species (Orthoptera: Caelifera)

The peering-jump behavior was studied for the common field grasshopper Chorthippus brunneus , the meadow grasshopper C. parallelus and the alpine grasshopper Miramella alpina (Orthoptera, Caelifera). It was found that immediately before jumping M. alpina executes primarily unilateral object-related peering movements, with approximately twice the amplitude and velocity of the predominantly bilateral object-related peering movements of the other two species. Whereas M. alpina almost always jumped toward the black stripes in the experimental arena, the other species jumped toward both the black stripes and the white spaces between them. All three species preferred the same pattern of black stripes, which permitted them to view one black stripe frontally, with an additional black stripe to the left and right, in the lateral visual field. The similarities and differences in the peering-jump behavior of the three grasshopper species is discussed with regard to visual perception (parallax cues) and environmental adaptation.     

Dose-related upwind anemotaxis and movement up odour gradients in still air in the presence of methyl eugenol by the wild tobacco fly, Bactrocera cacuminata

The behaviour of Bactrocera cacuminata (Hering) in wind varied according to the concentration of methyl eugenol (0, 95, 327 and 500 μg m^?3, respectively). General locomotor activity (as measured by mean distance moved in 5 min, regardless of direction) was not significantly different in the first two treatments but was significantly lower in the others. Most flies in the fourth treatment did not move more than one body length. In the first two treatments, the rate and pattern of movement of most flies was basically similar, with walking in tortuous paths interspersed with short flights and usually no obvious bias in direction. However, 32% of flies in the second treatment did move in a biased direction, achieving upwind anemotaxis of at least 400 mm, but only 2–8% did so in the other conditions. Flies moved up a concentration gradient to a source of methyl eugenol in still air when released at a distance of 100, 150 or 200 mm. With one exception, no more than 40% did this within 3 min of release (whether or not the olfactory stimulus was augmented by a visual one). However, 77% responded when released 100 mm from a combined olfactory and visual stimulus. Visual augmentation of an olfactory stimulus may also be responsible for far fewer flies flying out of the vicinity at distances up to 150 mm, but not 200 mm.     

A simple vision-based algorithm for decision making in flying Drosophila

Animals must quickly recognize objects in their environment and act accordingly. Previous studies indicate that looming visual objects trigger avoidance reflexes in many species [1-5]; however, such reflexes operate over a close range and might not detect a threatening stimulus at a safe distance. We analyzed how fruit flies (Drosophila melanogaster) respond to simple visual stimuli both in free flight and in a tethered-flight simulator. Whereas Drosophila, like many other insects, are attracted toward long vertical objects [6-10], we found that smaller visual stimuli elicit not weak attraction but rather strong repulsion. Because aversion to small spots depends on the vertical size of a moving object, and not on looming, it can function at a much greater distance than expansion-dependent reflexes. The opposing responses to long stripes and small spots reflect a simple but effective object classification system. Attraction toward long stripes would lead flies toward vegetative perches or feeding sites, whereas repulsion from small spots would help them avoid aerial predators or collisions with other insects. The motion of flying Drosophila depends on a balance of these two systems, providing a foundation for studying the neural basis of behavioral choice in a genetic model organism.