Sugar feeding and fluid destination control in the phlebotomine sandfly Lutzomyia longipalpis (Diptera: Psychodidae)

Sandfly feeding behaviour and destination of coloured sugar meals in the gut of Lutzomyia longipalpis were investigated with particular attention to the role of the crop. Sandflies were able to ingest sugar from liquid drops, microcapillaries, a slice of pear and even sugar powder. In most cases the flies adopted a 'sugar feeding mode' with raised palps. As the fruit dried, flies of both sexes fed by piercing the tissue with the proboscis. All sugar-fed flies had a full crop plus a small amount of sugary fluid in the thoracic mid-gut, i.e. past the stomodaeal valve. Dissections of flies interrupted during feeding showed that the very first trace of sugar passed through the stomodaeal valve, but that the rest of the meal was diverted into the crop. This suggests that closure of the stomodaeal valve is initiated only after a small volume of sugar solution has passed through it.     

Prandial lactate infusion inhibits spontaneous feeding in rats

To investigate the acute effects of lactate on spontaneous feeding, we infused lactate in the hepatic portal vein (0.5, 1.0, and 1.5 mmol lactate/meal) or in the vena cava (1.0 and 1.5 mmol lactate/meal) of ad libitum-fed rats during their first spontaneous nocturnal meal. Infusions (5 min, 0.1 ml/min) were remotely controlled, and a computerized feeding system recorded meal patterns. In separate crossover tests, meal size decreased independent of the infusion route after 1.0 and 1.5 mmol but not after 0.5 mmol lactate. The subsequent intermeal interval (IMI) tended to decrease only after vena cava infusion of 1.0 mmol lactate. The size of the second nocturnal meal increased after the 1.0 mmol lactate infusion. Hepatic portal infusion of 1.5 mmol lactate increased the satiety ratio [subsequent IMI (min)/meal size (g)] by 175%, which was higher than the insignificant 43% increase after vena cava infusion. Hepatic portal infusion of 1.5 mmol lactate also increased systemic plasma lactate but not glucose concentration at 1 min after the end of infusion. The results are consistent with the idea that meal-induced increases in circulating lactate play a role in the control of meal size (satiation). Moreover, the results suggest that lactate also contributes to postprandial satiety and that the liver is involved in this effect. The exact mechanisms of lactate's inhibitory effects on feeding and the site(s) where lactate acts to terminate meals remain to be identified.     

Feeding behaviour,midgut distension and ovarian development in Phlebotomus papatasi (Diptera: Psychodidae)

Phlebotomus papatasi females were fed through membranes or from cotton wool soaked in blood, water, sucrose or sodium chloride solutions. In membrane-fed flies, all diets were routed to the midgut and not to the crop. Following the meals that went to the midgut, females showed ovarian development at least 3 times greater than in sucrose-fed, autogenous control flies. Neither small quantities of water arriving in the midgut following drinking from soaked cotton wool, nor piercing of a membrane without feeding, stimulated ovarian development. Flies exhibited different feeding behaviour namely, blood feeding, sugar feeding, and water drinking. The blood-feeding behaviour was typical of flies ingesting any of the experimental diets through membranes, or blood or saline from cotton wool. The other two types of behaviour were observed in flies which fed from soaked cotton wool. The type of behaviour was characterized by the depth of penetration of the mouthparts into the substrate, the deployment of the palps and the degree of contact between the palps and the surface. It is suggested that the stimuli which control the routing of meals to the crop or to the midgut are derived from these types of behaviour.     

Post-feeding bubbling behaviour in fluid-feeding Diptera: concentration of crop contents by oral evaporation of excess water

Abstract During long post-feeding periods, adult tephritid flies (and apparently other non-blood feeding Diptera and even Hymenoptera) engage in behaviour consisting of oral extrusion and aeration of liquid droplets of varying size (‘bubbling’), proboscis pumping, and occasional deposition of regurgitate on the substrate that is followed by subsequent re-ingestion. Unlike evaporative cooling, which occurs in some insects when heat-stressed, these regurgitation behaviours take place regularly at moderate temperatures. To our knowledge, the significance of these behaviours has not been studied. We suggest that through bubbling behaviour, fully gorged Rhagoletis pomonella (Walsh) (Diptera: Tephritidae) flies eliminate excess water by evaporation to concentrate nutrients suspended in dilute solution, and release crop volume to allow resumption of fluid feeding or minimize the water load while foraging for other resources. Fly weights were measured continuously during pre- and post-feeding periods and in relation to occurrence of regurgitation behaviours. Fly weight losses during pre-feeding were an order of magnitude lower than post-feeding weight losses when flies exposed regurgitated liquid crop contents to air. During a bout of droplet extrusions, lasting on average 23 min, weight loss averaged 66% of the weight of liquid ingested by a fly in the preceding meal. Fly weight loss while bubbling was significantly correlated with duration of bubbling, temperature and relative humidity during post-feeding, and initial fly weight (adj. R² = 0.95). Fly age, volume of liquid ingested and rate of pre-feeding weight loss did not significantly improve predicted weight loss through bubbling. Findings are discussed in relation to other behaviours adopted by fluid-feeding insects to eliminate excess water.     

A model for blood meal digestion and fat metabolism in male tsetse flies (Glossinidae)

Abstract. Fat and haematin levels of mature male Glossina morsitans morsitans Westwood were estimated at different times after feeding at temperatures between 15 and 30°C. Flies were kept (largely inactive) in 7.5 × 2.5 cm tubes, or in actograph cages, where flight activity increased with time after feeding. Haematin excretion was modelled as a series of three first order reactions, all with the same rate parameter. The model accounted for > 98% of the variance in mean haematin in each of seven experiments; the rate parameter increased linearly with temperature and activity level. A similar approach was adopted for modelling fat metabolism. The rate coefficients of lipogenesis increased with temperature, and that for lipolysis with temperature, activity level and their interaction. All experiments were analysed simultaneously to provide equations predicting haematin or fat levels for all times, for active or inactive flies, and for temperatures between 15 and 30°C. Haematin exhibited large variations between individuals, but for active flies the expected haematin content at a given time varied little between flies kept at 25 and at 30°C. In inactive flies kept at 25°C, lipogenesis peaked at ≈ 24 h and lipolysis at ≈ 48 h. For active flies the times were 12 and 24 h, respectively; both rates were about twice as high as in inactive flies. Active flies produced (up to 1 mg) more fat out of a given size of blood meal than inactive flies. Curves of fat content against logarithm of haematin content differed little with temperature, and can therefore be useful for comparative studies of field populations of tsetse.     

Reproductive under-performance of tsetse flies in the laboratory, related to feeding frequency

Abstract. The rates of development of the eggs and larvae in utero and the next two developing ovarioles were measured by ovarian dissection on each day of the pregnancy cycle in tsetse, Glossina morsitans, subject to different feeding regimes. Compared with flies fed four times per pregnancy cycle, flies fed three times per cycle showed a lower pupal production rate (70%), the same (zero) adult mortality, a slightly slower growth rate of the larva and second ovariole only from day 8 onwards, but the same growth rate of the first ovariole. Flies fed only twice per pregnancy cycle produced no pupae, suffered 18% adult mortality and showed a significantly slower growth rate of the larva and second ovariole from days 6 and 7 respectively, but still the growth rate of the first ovariole was barely affected. Flies offered food three times or twice per pregnancy cycle engorged fully at every opportunity, but 16.5% of the flies offered food four times per cycle did not feed on every occasion, while 12–22% did not engorge fully on days 3, 5 or 7. In assessing the applicability of these laboratory results to the field situation the following points must be borne in mind: in the laboratory flies take smaller mean blood meals than in the field; during protein production associated with larval growth the proportion of the blood meal lost to transformation and excretory costs is less than during normal lipid metabolism; the balance between the tsetse's known fertility rate and adult and pupal mortality rates reveals that the abortion rate in the field must be extremely low. The high abortion rates usually observed in laboratory colonies, even when flies are offered food daily_l would be quite untenable in the field and indicate that laboratory conditions impose physiological stresses on the flies that are quite different from those in the field. These facts indicate that three field-sized meals may be sufficient to meet the energy demands of normal larval development in the field.     

Taste responses of Drosophila melanogaster

The amounts of sugar solution consumed by Drosophila melanogaster flies were determined. Starved and desiccated flies of a wild type strain (QA) consume 7?9 × 10^?2 λ of a 0.3 M sucrose solution per fly during the first hour and less later. They consume more of the 0.3 M sucrose solution than of the more diluted and the more concentrated solutions. In preference-aversion tests the flies discriminated between water and various sugar solutions, and between different sugar concentrations. Contrary to other fly species these flies did not prefer 0.05 M fructose over 0.05 M glucose. 0.3–0.5 M NaCl added to 0.1 M sucrose turned a preference over 0.01 sucrose into an aversion. A mutant, Lot-94, selected for its increased consumption of a 1 M NaCl solution was found to consume more of all test solutions. The amount of NaCl that had to be added to 0.1 M sucrose to turn the preference over 0.01 M sucrose by the mutant flies into aversion was not different from that found for the wild type flies.     

Micro-CT visualization of a promastigote secretory gel (PSG) and parasite plug in the digestive tract of the sand fly Lutzomyia longipalpis infected with Leishmania mexicana

Leishmaniasis is a debilitating disease of the tropics, subtropics and southern Europe caused by Leishmania parasites that are transmitted during blood feeding by phlebotomine sand flies (Diptera: Psychodidae). Using non-invasive micro-computed tomography, we were able to visualize the impact of the laboratory model infection of Lutzomyia longipalpis with Leishmania mexicana and its response to a second blood meal. For the first time we were able to show in 3D the plug of promastigote secretory gel (PSG) and parasites in the distended midgut of whole infected sand flies and measure its volume in relation to that of the midgut. We were also able to measure the degree of opening of the stomodeal valve and demonstrate the extension of the PSG and parasites into the pharynx. Although our pilot study could only examine a few flies, it supports the hypothesis that a second, non-infected, blood meal enhances parasite transmission as we showed that the thoracic PSG-parasite plug in infected flies after a second blood meal was, on average, more than twice the volume of the plug in infected flies that did not have a second blood meal.     

Volume of Protein meals taken by females of the blowfly, Lucilia cuprina: ovarian development-related and direct effects of protein ingestion

Abstract. The effect of the stage of ovarian development on the volumes of meals of the protein-rich material, liver exudate, taken by females of the blowfly, L. cuprina (Wiedemann), was examined using an experimental regime which was designed to isolate the effects of ovarian development from any other more direct effects of protein feeding or deprivation. It was found that volumes of meals taken by females with developing ovaries were little, if any, smaller than the volumes taken by protein-deprived females until quite late in vitellogenesis, at which stage meal size fell abruptly to that of meals taken by gravid females. A direct effect of protein-feeding was also demonstrated, with females that had consumed exudate during the previous few hours taking substantially smaller meals than those that had not done so. This effect was, however, shown to be non-specific in that the recent consumption of an equal volume of concentrated sucrose solution caused a similar reduction in the volume of exudate taken.     

Lifespan and patterns of accumulation and mobilization of nutrients in the sugar-fed phorid fly, Pseudacteon tricuspis

Abstract. The effect of sugar feeding on the survival of adult phorid fly Pseudacteon tricuspis is investigated. Flies fed 25% sucrose in aqueous solution continuously throughout their lifespan have greater longevity (mean ± SE longevity: female = 7.9 ± 0.8 days, male = 8.9 ± 0.9 days) than completely starved (provided no water and no sugar solution) flies, sugar-starved (provided water only) flies, or flies fed sugar solution only on their first day of adult life. Completely starved flies rarely lived beyond one day. Provision of water increases longevity by 2 days, and one full day of sugar feeding further increases longevity by an additional 1–2 days. Flies fed 50% sucrose have similar survivorship as those fed 25% sucrose. The temporal patterns of nutrient accumulation and utilization are also compared in P. tricuspis fed different diets: sugar-starved, sucrose-fed on the first day of adult life only, and sucrose-fed continuously. Adult P. tricuspis emerge with no gut sugars, and only minimal amounts of body sugars and glycogen. Although the levels of body sugars and glycogen decline gradually in sugar-starved flies, a single day of sugar feeding results in the accumulation of maximum amounts of gut sugars, body sugars and glycogen. High levels of these nutrients are maintained in female and male phorid flies fed on sucrose continuously over the observation period, whereas nutrient levels decline in flies fed only on the first day of life, beginning 1 day postfeeding. Female and male P. tricuspis emerge with an estimated 12.3 ± 2.3 and 7.2 ± 1 g of lipid reserves per fly, respectively. These teneral amounts represent the highest lipid levels detected in adult flies, irrespective of their diet, and are maintained over the life times of sucrose-fed female and male flies, but declined steadily in sugar-starved females. These data suggest that adult P. tricuspis are capable of converting dietary sucrose to body sugars and glycogen, but not lipids.     

Effects of substituted phenylethylamines on blowfly feeding behavior

The pharmacology of adult Phormia regina (Meigen) feeding behavior was explored by injecting candidate drugs into starved blowflies and then determining their responsiveness to aqueous sucrose, via the proboscis extension reflex. d-Amphetamine caused responsiveness to fall dramatically, while related drugs and biogenic amines had varying effects. When d-amphetamine-treated flies were fed 1 M sucrose, they consumed significantly more than control flies. Electrophysiological studies demonstrated that the responses of tarsal sugar receptor neurons to aqueous sucrose were not significantly altered by d-amphetamine. These observations are compatible with our hypothesis that octopamine positively modulates blowfly feeding behavior and suggest that other aromatic biogenic amines affect feeding behavior in this insect.     

Physiology of feeding-preference patterns of female black blowflies (Phormia regina Meigen): Alterations in responsiveness to salts

Electrophysiological recordings of tarsal and labellar contact chemosensilla in Phormia regina females demonstrate activation of sugar-, cation-, and possibly anion-sensitive cells in response to 10% yeast extract. Yeast extract is a proteinaceous food source that supports full egg development. Though some protein is required for egg maturation in Phormia, a blowfly not normally autogenous, purified proteins alone neither activate the chemosensilla nor promote egg development. In experiments reported here, groups of females were given ad lib access to either 0.2 M sucrose or 0.2 M sucrose and 10% yeast for four days after eclosion. The sucrose diet induced a ‘protein’ preference but no egg development. Potassium and sodium salts were added to 0.2 M sucrose and presented to the flies on day six. The volumes of salt-sucrose mixtures taken were greater than those of 0.2 M sucrose for flies raised on 0.2 M sucrose, though yeast was preferred over all but one salt solution. For sucrose-and-yeast-fed flies, volumes of 0.05 and 0.1 M salt mixtures consumed did not differ from 0.2 M sucrose; however, sucrose alone was favoured over more concentrated salt mixtures. All salt-sucrose mixtures tested were significantly more effective stimuli for sucrose-and-yeast-fed flies than yeast. It is suggested that varying sensitivity in the salt-sensitive contact chemosensilla plays a role in reported food preference shifts associated with reproduction.     

Post-alighting responses of Mexican fruit flies (Dipt., Tephritidae) to different insecticides in paint on attractive spheres

Abstract: Two new, comparatively safe insecticides (spinosad and imidacloprid) were compared with dimethoate (each at 1.5% active ingredient) for behavioural and mortality effects on Mexican fruit flies, Anastrepha ludens . Insecticide was mixed with sugar (as a feeding stimulant) and yellow latex paint (as an extending agent) applied to the surface of fruit-mimicking biodegradable 7 cm spheres made of sugar, flour and glycerin. Flies feeding on spinosad-treated spheres did not differ from flies feeding on untreated spheres in post-feeding intra-tree flight capability, amount of oviposition or mortality. Flies that fed on imidacloprid- or dimethoate-treated spheres for as little as 30 s experienced both high reduction in oviposition and high mortality compared with flies that fed on untreated spheres, and the flies from imidacloprid-treated spheres also showed a much reduced intra-tree flight capability. If baited with attractive odour, biodegradable yellow spheres treated with a surface coating of imidacloprid in latex paint and sugar could have potential for suppressing Mexican fruit flies on host trees.     

Stable fly, house fly (Diptera: Muscidae), and other nuisance fly development in poultry litter associated with horticultural crop production

Poultry litter usage in horticultural crop production is a contributor to nuisance fly populations, in particular stable flies (Stomoxys calcitrans L.) and house flies (Musca domestica L.). Extrapolation of adult emergence data suggests that approximately 1.5 million house flies and 0.2 million stable flies are emerging on average from every hectare of poultry litter applied as a preplant fertilizer for vegetable production in Perth, Western Australia. To a lesser extent, sideband applications to established crops may allow for the development of 0.5 million house flies and 45,000 stable flies per hectare. However, up to 1 million house flies, 0.45 million lesser house flies, Fannia cannicularis L., and 11,000 stable flies per hectare may be produced from surface dressings of poultry litter associated with turf production. Other nuisance flies present in poultry litter included the false stable fly, Muscina stabulans (Fallén), bluebodied blowfly, Calliphora dubia Hardy, black carrion fly, Hydrotaea rostrata Robineau-Desvoidy, Australian sheep blowfly, Lucilia cuprina Wiedemann, and flesh flies (Sarcophagidae). Only house flies developed in poultry litter for the first 4 d after application in the field. Stable flies were not present in poultry litter until 4-7 d after application, and were the only fly species developing in litter > 9 d after application.     

Physiology of feeding preference patterns of female black blowflies (Phormia regina Meigen)

Summary The relative effectiveness of 0. 1 M sucrose and 10% yeast extract in eliciting feeding by femalePhormia regina has been shown to be related to the fly's carbohydrate reserve levels. Females that have had ad lib. access to 0.1 M sucrose (S flies, standard condition) take larger drinks of 10% yeast than 0.1 M sucrose starting at three days of age, indicating that 10% is the more effective feeding stimulus. Females that have had ad lib. access to 0.1 M sucrose and 10% yeast (S/Y flies, standard condition) always drink more 0.1 M sucrose than 10% yeast (Fig. 1). S/Y (standard condition) flies appear to respond to the yeast as if it were dilute carbohydrate: the volumes of 0.1 M sucrose and 10% yeast that they drink fluctuate in parallel. S (standard condition) flies respond to the yeast differently: starting day 3, their drink volumes of 10% yeast increase as drink volumes of 0.1 M sucrose decrease.The elevation of the volume of 0.1 M sucrose drunk by flies whose carbohydrate reserve levels have been depleted by diet, starvation or flight (Figs. 2–4) supports the hypothesis that some physiological state or process related to carbohydrate metabolism regulates the fly's responsiveness to yeast extract. Possible mechanism by which the fly translates carbohydrate reserve level into feeding preference are discussed. Available evidence points to a CNS phenomenon, probably involving the stomatogastric nervous system. The pattern of the selective feeding cycle of femalePhormia can be explained in terms of the accumulation and utilization of carbohydrate reserves.This work was supported by a Spencer Foundation Predoctoral Fellowship from Princeton University and by NSF Grant number BNS 76-02172 to Dr. Vincent G. Dethier. I am grateful to the Departments of Zoology and Entomology of the University of Massachusetts, Amherst, for the use of their facilities. Many thanks to Drs. V.G. Dethier, Elizabeth Bowdan and David Falk for critical discussion and assistance.     

Mushroom host influence on Lycoriella mali (Diptera: Sciaridae) life cycle

Lycoriella mali Fitch (Diptera: Sciaridae) infests mushroom crops early in the crop cycle. Recent observations in mushroom houses indicated a difference in emergence time and size of adult L. mali developing on various strains of commercial mushrooms. Samples of adult flies from isolated mushroom houses growing Portabella mushrooms were significantly heavier then those from oyster mushroom houses, whereas flies from shiitake mushroom houses were lightest in weight. Flies collected from isolated Portabella mushroom houses were reared on four strains and species of Agaricus and Pleurotus mushrooms. After the adults emerged, females were weighed, mated, and allowed to oviposit. The number of eggs laid increased as the weight of the female increased. Flies collected from isolated Portabella mushroom houses were reared on eight strains and species of mushrooms. Flies were reared for four generations on each host mushroom mycelium then switched to different host mushrooms. Overall, the hybrid strain of Agaricus bisporus (Lange) Imbach (Agaricales: Agaricomycetideae) was the most favorable host for L. mali, whereas the wild strain of A. bisporus was the least favorable host. Mushroom hosts influence developmental time, survivorship, weight, and reproduction of L. mali.     

Structural evaluation of anorectic action induced by 1,5-anhydro-D-glucitol

In previous studies, 1,5-anhydro-D-glucitol (1-DG), an endogenous glucose analog, was found to significantly influence physiological feeding behavior. The relationships between the hydroxyl group positions on the pyranose ring carbons and the anorectic action induced by 1-DG and its analogs are discussed. To investigate the effects of these glucose analogs on ingestive behavior, 24 mumole of test solution was injected into the rat third cerebral ventricle immediately before normal eating time, which starts at the beginning of the dark. After initial transient hyperphagia, 1-DG suppressed feeding during the first 12-hr dark period. It prolonged postprandial intermeal interval beginning shortly after injection, but eating rate was not affected and meal size did not decrease until near the end of the normal feeding period. The incidence of drinking episodes decreased concomitant with feeding suppression. Feeding and drinking suppression were also produced by 1,2-dideoxy-D-glucose, 1,3-dideoxy-D-glucose, and 1,4-dideoxy-D-glucose, although they were less potent than 1-DG. They suppressed feeding by prolonging the postprandial intermeal interval, but did not change meal size or eating rate. The anorectic effects of 1-DG were abolished by removal of the hydroxyl group at carbon 6 and by epimerization at carbons 2, 3, and 4. These findings indicate that feeding suppression induced by 1-DG and its analogs is induced mainly by prolongation of the postprandial intermeal interval, and the presence or absence of a hydroxyl group on each carbon of 1-DG is important for its feeding suppression.     

Some effects of light and heat on the feeding and resting behaviour of tsetse flies,Glossina morsitans Westwood

A laboratory investigation was carried out into the effects of light and heat on the feeding and resting behaviour of Glossina morsitans morsitans Westwood. Flies placed in small cages on the ears of rabbits were tested for feeding responsiveness at light intensities of 0, 10, 100, 1000 and 25 000 lux. Feeding occurred at the same rate in darkness as it did in light at up to 1000 lux, but at 25 000 lux was largely inhibited. At 26° C, flies flew to and settled in the light end of a 1.2 m-long choice chamber. As the temperature was raised (by 10° C/hour) they eventually took off and flew directly to the dark end. The temperature at which this skototaxis occurred was strongly negatively correlated with the light intensity: each log unit increase in light intensity (in lux) caused a 2.2° C fall in take-off temperature. Skototaxis occurred at higher temperatures in recently fed female flies than in starved ones.     

Transmission of MdSGHV among adult house flies, Musca domestica (Diptera: Muscidae), occurs via oral secretions and excreta

The MdSGHV is a double-stranded DNA virus that replicates in the salivary glands of infected adult house flies. Transmission of this non-occluded, enveloped virus is believed to be mediated orally via deposition and consumption of oral secretions composed of salivary gland secretions and crop contents. In this study, transmission electron micrographs of crops from infected flies showed numerous enveloped virions in the crop lumen adjacent to the cuticular intima, as well as on the hemocoel side in close vicinity to muscle cells. Oral treatments of newly emerged flies with viremic salivary gland homogenates, crop homogenates, or gradient-purified virus resulted in an average 44% infection. Virus released via oral secretion was infectious when ingested by newly emerged adult flies, resulting in an average 66% infection. Using quantitative real-time PCR, MdSGHV DNA was quantified in oral secretions and excreta obtained from viremic flies. Between 2 and 4 days post-infection (dpi), viral copy numbers in oral secretions increased exponentially and from 5 to 21 dpi each infected fly released an average 10⁶ MdSGHV copies per feeding event. Excreta samples collected overnight from individual infected flies at 5 dpi contained an average 6.5 × 10⁵ viral copies. Low but detectable infection rates were produced when newly emerged flies were challenged with excreta samples. In summary, evaluation of the quantity and infectivity of MdSGHV released by individual infected house flies clearly showed that deposition of oral secretions and excreta onto a shared food substrate is the main route of natural MdSGHV transmission among adult house flies.     

Temporal patterning of feeding by hummingbirds

For five species of hummingbirds in the laboratory, time between meals was related to energy intake on the first meal and rate of energy expenditure between meals. Field observations gave similar results. Average meal sizes were similar at one intake rate independent of food caloric density; females averaged longer bouts than males. When rate of intake was approximately halved, meal duration approximately doubled and volume intake remained similar. We postulate that feeding is initiated when crop contents reach a lower threshold and that feeding is terminated after ingestion of an optimal volume determined by the added weight of the meal.