A theory of the pattern induced flight orientation of the fly Musca domestica

The theory presented here describes the visual orientation behavior of fixed flying insects (the fly Musca domestica) in the presence of elementary patterns. The theory, which is based on a number of experimental results, Reichardt (1973), is a phenomenological one whose main purpose is to provide an organizational framework for treating a complex phenomenon without the need of detailed assumptions about the neural mechanisms actually involved.A simple hypothesis concerning the basic structure of the pattern fixation process leads to an equivalent stochastic equation of the Langevin type, which can be linearized for simple single-stripe panoramas. A critical experiment supports these theoretical assumptions. In addition, the effect on pattern fixation behavior of adding contrast noise to the background of the panorama, is quantitatively predicted by the theory.In the more general case of a panorama consisting of many vertical stripes, the Fokker-Planck equation associated with the Langevin equation, no longer linear, is solved. Making use of an experimentally proven superposition principle, the stationary pattern fixation behavior of the fly in an arbitrary panorama consisting of a collection of vertical stripes is predicted. In this context, concepts like pseudo-invariance and phase-transition can be applied to the insects orientation behavior. The theory presented here seems to contain rich classification properties, which might provide the foundations for an understanding of more complex pattern discrimination processes.Possible extensions of the theory, as well as some similarities to human eye fixation, are also discussed.     

Visually induced height orientation of the fly Musca domestica

The visually controlled height orientation of fixed flying flies (Musca domestica) was investigated. The flight lift force measured by a transducer drives the vertical motion of a panorama. The dynamical conditions of the free flight are electronically simulated for the fly with respect to this degree of freedom of motion. In most of the experimentally investigated cases the panorama consists of a horizontally oriented narrow dark stripe on a bright background. The fly orientates with respect to the stripe, transporting it into a stable fixation position just below the equatorial plane of its compound eyes. It is experimentally demonstrated that the formalism of the linearized theory of the pattern induced flight orientation — Poggio and Reichardt (1973a) — can be applied to describe the height orientation of the fly. The experimental evidence concerning the simultaneous perception of stripes moving in a well defined manner in front of each of the two compound eyes is consistent with the hypothesis that the two halves of the visual system are perceptually additive.     

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.     

Ecdysteroidostatin from the house fly,Musca domestica

Ovaries from house flies maintained on sucrose secrete large amounts of ecdysteroid when they are cultured with ovarian ecdysteroidogenic hormone, OEH. However, ovarian ecdysteroid secretion is reduced by incubation with both OEH and the ovarian ecdysteroidostatin (OES). A partially purified OES fraction from a semi-preparative reverse phase HPLC C₁₈ column caused a 98% inhibition of ovarian ecdysteroid secretion in vitro at a concentration of 0.8 equivalents per μl. Ovaries can be activated to produce ecdysteroid in vivo by feeding diet containing protein to flies maintained on sucrose. Ecdysteroid secretion was inhibited when the in vivo stimulated ovaries were cultured with OES. This suggests that OES does not interfere with the OEH activation mechanism, but blocks ovarian ecdysteroid synthesis or release. Furthermore, OES inhibition is reversible and ecdysteroid secretion resumes when OES is removed. Musca OES could explain the decrease in ecdysteroid levels found in flies after mid-vitellogenesis. Both adult male and female abdomens contain OES, but OES was not transferred to females during mating. Evidence is presented that OES is not a trypsin modulating oostatic factor. Arch. Insect Biochem. Physiol. 38:166–176, 1998. © 1998 Wiley-Liss, Inc.     

Asymptotic oscillations in the tracking behaviour of the fly Musca domestica

From recent theoretical work (Poggio and Reichardt, 1981), high frequency oscillations are expected in the angular trajectory of houseflies tracking a moving target if the target's retinal position controls the flight torque by means of a stronger optomotor response to progressive than to regressive motion. Experiments designed to test this conjecture have shown that (a) asymptotic non-decaying oscillations are found in the torque of female houseflies tracking targets moving at constant angular velocity; (b) the magnitude of the oscillations grows monotonically with mean retinal excentricity of the target; (c) the period of the oscillation is around 180–200 ms. The experimental findings are consistent with the hypothesis that a progressive-regressive mechanism plays a significant role in the tracking behaviour of female houseflies. From this phenomenological point of view a flicker mechanism that is active only for nonzero motion is equivalent to a progressive-regressive system. The relatively long period of the oscillation requires more complex reaction dynamics than a pure single dead-time delay. As a specific example we show that a model where the reaction to progressive motion is sticky, holding for a longish time after the ending of the stimulus, is consistent with the experimental data.     

Vibrio cholerae laboratory infection of the adult house fly Musca domestica

The present study was designed to test the hypothesis that house flies may be capable of specifically harbouring ingested Vibrio cholerae in their digestive tracts. Flies were continuously fed green fluorescent protein (GFP)‐labelled, non‐O1/non‐O139 environmental strains of V. cholerae. Bacterial burdens were quantitatively measured using plate counts and localization was directly observed using confocal microscopy. Vibrio cholerae were present in the fly alimentary canal after just 4 h, and reached a plateau of ～10⁷ colony‐forming units (CFU)/fly after 5 days in those flies most tolerant of the pathogen. However, individual flies were resistant to the pathogen: one or more flies were found to carry < 180 V. cholerae CFU at each time‐point examined. In flies carrying V. cholerae, the pathogen was predominantly localized to the midgut rather than the rectal space or crop. The proportion of house flies carrying V. cholerae in the midgut was dose‐dependent: the continuous ingestion of a concentrated, freshly prepared dose of V. cholerae increased the likelihood that fluorescent cells would be observed. However, V. cholerae may be a transient inhabitant of the house fly. This work represents the first demonstration that V. cholerae can inhabit the house fly midgut, and provides a platform for future studies of host, pathogen and environmental mediators of the successful colonization of this disease vector.     

Search orientation of Musca domestica in patches of sucrose drops

ABSTRACT. The searching tactics of the housefly, Musca dormestica L. (Diptera: Muscidae), have been delineated from digitized pathways of flies walking in patches of sucrose drops arranged in linear (ROW) and hexagonal (HEX) arrays. The areas covered by flies in ROW and HEX patches are distinctly different, but flies seem not to employ different tactics for the two types of resource arrays. The number of drops located, if at least one drop is found, does not differ between ROW and HEX. Most quantitative measures of local search remain constant after the first interdrop interval, although feeding time decreases as flies sample successive drops. Local search intensifies after each drop is ingested, with locomotory rate decreasing and turning rate increasing, followed by decay of both measures toward ranging levels. Searching can be characterized by two movement tendencies resulting from specific, definable, locomotory functions: a forward-moving tendency is expressed by the fly as it leaves a resource in approximately the same direction as it arrives; and local search is characterized by looping, rather than straight walking, with a variable turning rate that generates a rough ‘zigzag’ superimposed on looping. The two movement tendencies, combined, allow flies to locate resources in a linear arrangement, because of the forward-moving tendency, and to locate resources not arranged in a linear array because of the ‘noisy’ loop. M. domestica does not appear to retain and use information gained from one patch of drops in another, so the search tactic of the fly seems therefore to be a compromise between straight movement and circular movement that may be adaptive for an environment subject to frequent changes in the spatial distribution of resources. Giving-up-time, the period between ingesting the last drop and leaving the patch, is a function of the rate of change in the transition from local search to ranging, which is constant in our experiments. If a fly does not encounter another drop while ranging, during which it walks relatively straight, the fly moves out of and away from the patch.     

Genetic control of LDH isozymes in the house fly,Musca domestica

Electrophoretic variations in lactate dehydrogenase from adult whole body homogenates are described for three laboratory strains of house fly, Musca domestica. Several crosses between different electrophoretic forms provided evidence that the observed variations are due to segregation of alleles at two distinct loci (designated as A and B loci) and that the LDH isozymes of house flies are dimers formed by a random association of subunits controlled by the two loci.     

The cytochrome C oxidase of the house fly,Musca domestica L

1. Insoluble and "soluble" forms of cytochrome c oxidase were prepared from house flies by methods described for mammalian tissue. A 4.5-fold purification was accomplished by preparing the enzyme in insoluble form (as compared with a crude water brei). The oxidase in "soluble" form was purified 20.0-fold. 2. The absorption spectrum of the "soluble" preparation from flies was similar to that from mammalian tissue, and indicated the presence of cytochromes a + a(3) and b. In contrast with the corresponding mammalian enzymes these components were not separable by the existing techniques. 3. The Soret peak seen at 440 mmicro and the alpha peak at 600 mmicro were due. primarily to cytochromes a(3) and a, respectively. 4. The oxidase activity of whole house flies was comparable with that of rat abdominal muscle. In both organisms, this activity was inhibited by cyanide to approximately the same extent. 5. The optimum buffer concentration for oxidase activity, as measured spectrophotometrically, was approximately 0.02 to 0.06 M. This is in agreement with results obtained for mammalian enzyme, as measured both manometrically and spectrophotometrically. 6. The present study has demonstrated the following components of the cytochrome system in house flies: b, c, a, and a(3).     

Cuticular pheromones and water balance in the house fly,Musca domestica

Epicuticular lipids serve two major roles in insects. Their waterproofing properties are crucial to survival in terrestrial environments, and they serve as contact pheromones in a wide array of taxa. Both functions may be affected by the physical properties of the surface lipids. This provides the opportunity for natural selection on water conservation, mediated by lipid phase behavior, to interact with and perhaps conflict with sexual selection on communication and mate recognition. We used the common house fly, Musca domestica, as a model for these interacting selective forces. Male house flies preferred female models treated with a high melting-point lipid mixture, suggesting that sexual and natural selection may both act to favor longer-chain, more saturated hydrocarbons. However, higher melting points did not result in lower rates of water loss. We propose a working model in which phase separation between the unsaturated female pheromone and saturated hydrocarbons results in areas of melted, pheromone-rich lipids and regional variation in cuticular permeability.     

Disease dynamics and persistence of Musca domestica salivary gland hypertrophy virus infections in laboratory house fly (Musca domestica) populations

Past surveys of feral house fly populations have shown that Musca domestica salivary gland hypertrophy virus (MdSGHV) has a worldwide distribution, with an average prevalence varying between 0.5% and 10%. How this adult-specific virus persists in nature is unknown. In the present study, experiments were conducted to examine short-term transmission efficiency and long-term persistence of symptomatic MdSGHV infections in confined house fly populations. Average rates of disease transmission from virus-infected to healthy flies in small populations of 50 or 100 flies ranged from 3% to 24% and did not vary between three tested geographical strains that originated from different continents. Introduction of an initial proportion of 40% infected flies into fly populations did not result in epizootics. Instead, long-term observations demonstrated that MdSGHV infection levels declined over time, resulting in a 10% infection rate after passing through 10 filial generations. In all experiments, induced disease rates were significantly higher in male flies than in female flies and might be explained by male-specific behaviors that increased contact with viremic flies and/or virus-contaminated surfaces.     

Innate control of local search behaviour in the house fly, Musca domestica

Abstract. The genetic basis of variation in local searching behaviour in the house fly, Musca domestica L., was examined by estimating the heritability of locomotory parameters using parent—offspring regressions. Pathways of adult flies were videotaped for 3 min after ingestion of a 1 μl drop of 0.25 M sucrose in a 75 cm diameter circular arena with minimal external cues. Locomotory variables computed from digitized walking pathways, including locomotory rate, turning rates per unit of distance and time, turn bias, number of stops, and stop duration, were significantly heritable, but heritabilities (additive genetic variance ÷ total phenotypic variance) was generally low (<0.25). Phenotypic variance in search parameters increased with time since ingestion of the sucrose drop, but heritabilities were highest when only the first 30 s of the searching bout were examined, and declined as longer periods were considered. Significant genetic correlations among search parameters suggest that many of these characters are influenced pleiotropically by the same genes or closely linked genes. However, some aspects of local search were inherited independently (e.g. stop duration and locomotory rate). The independence of controlling mechanisms was supported by observed differences between genetic and environmental correlations. This genetic independence could enhance the likelihood of appropriate population responses to temporally or spatially changing selective forces.