The potential for control of the blowfly Lucilia sericata using odour-baited targets

Abstract A deterministic simulation model is used to explore the potential for control of the sheep blowfly, Lucilia sericata , using odour-baited targets. The simulations show that the ambient temperatures of 15^oC or 20^oC, 13% and 24% of the females present in a population would need to be killed per day, respectively, to prevent the population growing from its initial spring density during the summer blowfly season. In contrast, if both sexes could be sterilized, so that sterilized males were also able to disrupt the fertility of unsterilized females, only 6.5–13% of both sexes would need to be attracted and sterilized at 15^oC or 20^oC respectively. To examine whether these numbers of individuals could be attracted by targets baited with liver and sodium sulphide, mark-release-recapture studies in sheep pastures were used to quantify sticky target efficiencies. The percentage of the Lsericata population caught on any one day was shown to be positively related to average ambient temperature. At 15^oC each sticky target caught approximately 13% and at 20^oC, 41% of the females within a 20 m radius. The results indicate that, at ambient temperatures between 15 and 20^oC, if distributed at a density of approximately 5/ha, targets baited with liver and sodium sulphide could effectively suppress a population of L.sericata. Lower densities of targets would be required if they were able to sterilize both sexes rather than kill or if more attractive synthetic semiochemical baits could be developed.     

Effects of targets impregnated with the chitin synthesis inhibitor triflumuron on the blowfly Lucilia sericata

The effects of exposure of adults of the blowfly Lucilia sericata (Meigen) for 24 h to cloth targets impregnated with the chitin synthesis inhibitor triflumuron plus sucrose or sucrose only, were considered. When both male and female L. sericata were exposed to targets treated with sucrose only, the percentage egg hatch varied between 60–75%. When both sexes were exposed to targets treated with 10% triflumuron plus sucrose the percentage egg hatch varied between 3–7%. When females alone were exposed to triflumuron, prior to the introduction of unexposed males, egg hatch varied between 4–14%. The exposure of males alone to 10% triflumuron treated targets, prior to the introduction of unexposed virgin females, did not give any consistent significant decrease in the mean percentage egg hatch. However, repetition of this trial with males exposed to targets treated with 15 or 20% triflumuron, resulted in a percentage egg hatch of 3.1% and 10.3%, respectively. These results suggest that at relatively high target concentrations, males can transfer effective doses of triflumuron to unexposed virgin females.The exposure of adults to triflumuron treated targets did not significantly increase the mortality of either male or female L. sericata. Furthermore, bioassays showed that there was no significant decline in the ovicidal effectiveness of targets treated with 10% triflumuron and sucrose solution following exposure to sun and rain for up to 8 weeks in 1995 and 1996.The results suggest that targets treated with triflumuron suspension concentrate and sucrose may form the basis of an effective system for the control of L. sericata in the field. However, the relatively high concentrations of triflumuron required and the low efficiency with which males are able to transfer effective doses to unexposed females may diminish one of the main theoretical advantages of using a chitin synthesis inhibitor, such as triflumuron, over a conventional insecticide.     

The use of carrion as breeding sites by the blowfly Lucilia sericata and other Calliphoridae

Abstract. To examine the species composition of flies breeding in carrion in the field, the carcasses of mice and quail were exposed on sheep farms in the South West of England. Calliphora vicina was the dominant species of Diptera; 19,294 individuals emerged from 175 of the 241 infested carcasses recovered. Lucilia sericata emerged from only 39 of the infested carcasses, at a median of 10 adults per infested carcass. Other species of Lucilia present were L.ampullacea, L.caesar and L.illustris. The highest number of L.sericata emerged from carcasses placed in open pasture, the highest number of C.vicina emerged from carcasses in hedgerow, whereas the highest numbers of L.caesar, L.ampullacea and L.illustris emerged from carcasses in woodland. The duration of exposure of carcasses in the field was negatively related to the size of the adult L.sericata which emerged and, in woodland and hedgerow habitats, to the number of L.sericata which emerged. These data indicate that the larvae of L.sericata in carcasses experience significant levels of competition and that the intensity of this competition may be sufficient to reduce the numbers of L.sericata able to emerge successfully. The size distributions of female L. sericata which emerged from carcasses or which were caught as adults in the field showed only a small degree of overlap, suggesting that only a relatively small proportion of the wild L.sericata population emerge from carcass breeding sites. The results are discussed in relation to the development of new approaches to the control of blowfly strike of sheep.     

Responses of the sheep blowflies Lucilia sericata and Lcuprina to odour and the development of semiochemical baits

Abstract. The literature relating to the attraction of the sheep blowflies Lucilia sericata and Lucilia cuprina to their ovine hosts is reviewed. The responses of the two species are similar and different components of the behaviour leading to host location and oviposition appear to involve at least two distinct sets of semiochemical cues. Activation, upwind orientation and landing appear to occur in response to putrefactive sulphur-rich volatiles, originating from bacterial decomposition products. Oviposition is elicited primarily by the presence of ammonia-rich compounds; moisture, pheromones and tactile stimuli may also act as oviposition stimuli. There is a pronounced sex difference in the response of Lucilia to semiochemicals with a higher proportion of females attracted than males and a higher proportion of gravid than non-gravid females. While the mechanisms of host location by Lucilia are of intrinsic interest, understanding the responses to semiochemicals is important in the attempt to develop powerful synthetic baits for deployment with the traps or targets used for population sampling or suppression. The literature is discussed with respect to the development of synthetic semiochemical baits.     

A comparison of Lucitraps and sticky targets for sampling the blowfly Lucilia sericata

The Lucitrap (Miazma Pty Ltd, Queensland, Australia) combined with a synthetic odour bait, Lucilure (Miazma Pty Ltd, Queensland, Australia), is a commercially available trap for sampling and control of Lucilia cuprina (Wiedemann) in Australia. It was tested in Hungary against Lucilia sericata (Meigen) (Diptera: Calliphoridae), a cause of sheep strike throughout temperate Europe. The standard Lucitrap was tested against black or yellow sticky target traps. Both trap types were baited with either Lucilure or liver and 10% w/v sodium sulphide solution. With Lucilure as bait, L. sericata were caught on sticky traps but not in Lucitraps. With liver and sodium sulphide as bait, sticky traps caught 500-1500 times more L. sericata than Lucitraps. An adhesive sheet fitted to the top of a Lucitrap captured 30-300 times more L. sericata then were captured inside an unaltered Lucitrap. Direct observation of metallic green calliphorids (92.1% L. sericata) alighting on Lucitraps indicated that most flies stayed for a short while (modal class 2-4 s) and only a few stayed longer, to an observed maximum of 28 s. Flies explored a mean of 1.5 entry holes (range 0-7) during a visit but only 6% entered the trap. Size of L. sericata was not a physical barrier to Lucitrap entry, because many larger species were captured. However, L. sericata captured inside Lucitraps were significantly smaller than those captured on sticky traps, demonstrating that size was of behavioural importance. The data demonstrate that the Lucitrap is not effective as a trap for L. sericata in Hungary, due mainly to a failure of flies to enter the trap in large numbers. In Australia and South Africa, L. sericata is commonly caught in Lucitraps baited with Lucilure, although L. cuprina is more numerous. Our study highlights the potential for diversity of fly behaviour between different geographical populations of the same species. Such diversity can have a significant effect on the functioning of systems for fly sampling and control, when these systems depend for their success on certain behavioural responses of the target species.     

Pre-adult mortality in the blowfly Lucilia sericata

Populations of the blowfly, Lucilia sericata (Meigen) (Diptera: Calliphoridae), have a considerable potential for rapid increase; the lifetime reproductive output of each adult female has been estimated to be between 130 and 172 eggs. Nevertheless, in the field, absolute population densities of this species are relatively low. To account for this difference, the levels of mortality affecting the eggs, feeding and wandering larvae and pupae of L. sericata were assessed in the field and laboratory. Percentage egg hatch was dependent on relative humidity with no egg eclosion at humidities below 50%; there was no significant effect of temperature on egg hatch. On infested sheep, the mean mortality of feeding larvae was 53%, but this ranged widely from 0% to 96%. There was no effect of atmospheric temperature or humidity on the mortality of feeding larvae in vivo. In the laboratory, only 10% of wandering larvae pupariated at 10 degrees C. At above 20 degrees C pupariation was consistently almost 100%. Percentage emergence increased from 0% at 10 degrees C to about 80% between 20 and 30 degrees C. The upper lethal temperature for pupae was approached at 35 degrees C. Analysis of the predation of pupae in the field revealed a weak, but significant curvilinear relationship between temperature and proportionate mortality and a median mortality of 0.49% per 24 h exposure (interquartile range = 5.2%). There was no evidence of density dependence in pupal predation. Overall, it is estimated that pre-adult mortality accounts for losses of approximately 97% of each generation, but this figure is subject to considerable variation depending on factors such as climate, time of year and host susceptibility.     

Visual and olfactory cue interaction in resource-location by the blowfly, Lucilia sericata

Abstract. The interactions between olfactory and visual cues in the landing responses of the blowfly Lucilia sericata (Meigen) (Diptera: Calliphoridae) were examined in a laboratory wind tunnel. The odour of liver and sodium sulphide solution released at 1 L/min, resulted in a greater number of landings, closer to, but downwind of, the central odour release point, than when odour was absent. Three-day-old protein-deprived females landed in greater numbers than protein-fed female flies of the same age; no difference existed between 6-day-old protein-fed or protein-deprived females. Six-day-old, protein-fed flies landed closer to the odour source than did 3-day-old, protein-fed flies. In the presence of odour, flies landed closer to the central release point when it was accompanied by a visual cue. No such effect of the visual cue was evident in the absence of odour. When a plain white sticky-surface (25 × 25 cm) was paired with an identical white surface plus odour, approximately equal numbers of flies landed when the surfaces were placed together; increasingly higher numbers of flies landed on the surface with the odour cue when the distance between the surfaces was increased. When a white surface with a visual cue was presented with the plain white surface plus odour, more flies landed on the white surface with the visual cue than on the plain white surface with odour when they were close together. However, as these two surfaces were moved apart, flies landed increasingly more on the surface with the odour than the surface with the visual cue. The results show that while odour cues may enhance the induction of landing by female L. sericata , visual cues are important when selecting a final landing site.     

Colour discrimination by the sheep blowfly Lucilia sericata

Abstract. In laboratory trials designed to examine the alighting response of the blowfly Lucilia sericata Meigen to colour, yellow sticky cloth targets caught the largest number of both males and females, followed by pale blue, black, green, dark blue and red. The number caught by any colour showed a strong positive relationship with the percentage of its spectral reflectivity in the 450–580 nm wavelength band. In field trials, targets baited with the synthetic attractant 'swormlure' caught significantly fewer L.sericata than targets baited with liver and sodium sulphide, suggesting that the former bait is a relatively poor attractant for this species, at the release rates used in the present study. However, there was no effect of target colour on catch, neither was there any interaction between colour and odour bait type. The results, from both the laboratory and field trials, suggest that the strength of responses by Lsericata to visual cues are weak relative to responses to odour. Responses to hue are readily overridden by a range of factors such as brightness and contrast with the background and are therefore likely to be difficult to detect or manipulate reliably in trapping systems in the field.     

Suppression of the blowfly Lucilia sericata using odour-baited triflumuron-impregnated targets

Abstract. Field trials were carried out in 1995 and 1996 on farms in the south-west of England to assess the extent to which odour-baited targets could be used to suppress populations of the ectoparasitic blowfly, Lucilia sericata , in sheep pastures. Targets were constructed from 41 × 41 cm squares of aluminium sheet, covered by white cloth which had been dipped in a mixture of sucrose solution (50% w/v) and the chitin synthesis inhibitor triflumuron (10% suspension concentrate). Each target was baited with ≈ 300 g of liver and sodium sulphide solution (10%). Three matched sheep farms were used in the trials. In 1995, triflumuron-impregnated targets were placed around the periphery of sheep pastures at one of the farms in late June, at approximately one target per hectare. In 1996, triflumuron-impregnated targets were placed around the periphery of sheep pastures of a second of the farms in early May, at approximately five targets per hectare. Each year, five sticky targets, used to monitor the L. sericata populations, were also placed in fields at the experimental and the other two farms, which acted as controls. In 1995, the results provided some, although inconclusive, evidence that the triflumuron-impregnated targets had reduced the numbers of L. sericata relative to the populations on the two control farms. In 1996, however, the density of L. sericata on the experimental farm was reduced to almost zero and remained significantly lower than on two control farms throughout the period during which the triflumuron-impregnated targets remained in the field. The results are discussed in relation to the use of triflumuron-treated targets as a practical means of controlling L. sericata and sheep blowfly strike.     

Mortality rate, reproductive output, and trap response bias in populations of the blowfly Lucilia sericata

1. The ages of 877 females of the blowfly Lucilia sericata, collected from two farms in south-west England between May and September 1996, were determined using a combination of ovarian dissection and wing fray analysis. 2. Using survivorship analysis, the mortality rates over the entire field season were estimated to be 2.2 and 1.9% per day-degree at the two farms, respectively. These gave a mean life expectancy of 46 and 53 day-degrees, and a lifetime reproductive output of 130 and 172 eggs per female. 3. The mortality rate remained relatively constant over time, with one exception, the first sample of the season when no older flies were present, indicating that underlying changes in demography (emergence and senescence of generations) over the field season were not sufficient to affect the rates calculated by survivorship analysis. 4. The response of L. sericata to liver-baited traps was affected strongly by age and reproductive status, with gravid females and females in the early stages of ovarian development being most markedly over-represented relative to other age categories.     

A simple larval diet for population studies on the blowfly Lucilia sericata (Diptera: Calliphoridae)

1. A simple artificial diet was devised for larvae of Lucilia sericata Meigen. 2. A basic diet of 20 g/l agar with 20% horse blood and added yeast sustained normal growth and development, as compared with a lamb meat control. 3. When yeast was added to the blood agar diet at 50 g/l, both peak and final larval weights were increased by 25-50% at higher larval densities (2-8 larvae/g diet) and the hatchling-adult development period was reduced by about 3 days. 4. No adult insects emerged on an agar-yeast diet without blood. Increasing the concentration of blood from 10% to 20% increased adult weight at emergence by 50%, though developmental period was not significantly affected. 5. Increasing larval density significantly reduced the weights of pupae and emergent adults (by up to 50%), both on the blood-agar-yeast diet and the lamb meat control. Percentage survival fell from 80-90% to below 10% on the blood-agar-yeast diet at the highest densities of 4 or 8 larvae/g diet, though survival on lamb was not significantly affected by density.     

Modelling the impact of climate change on spatial patterns of disease risk: sheep blowfly strike by Lucilia sericata in Great Britain

Understanding the spatial scale and temporal pattern of disease incidence is a fundamental prerequisite for the development of appropriate management and intervention strategies. It is particularly critical, given the need to understand the elevated risks linked to climate change, to allow the most likely changes in the distribution of parasites and disease vectors to be predicted under a range of climate change scenarios. Using statistical models, the spatial distribution and climatic correlates of a range of parasites and diseases have been mapped previously, but their development into dynamic, predictive tools is less common. The aim of the work described here, was to use a species distribution model to characterise the environmental determinants of the monthly occurrence of ovine cutaneous myiasis (blowfly strike) by Lucilia sericata, the most frequent primary agent of northern European myiasis, and to then use this model to describe the potential spatial changes that might be expected in response to predicted climate change in Great Britain. The model predicts that the range of elevated temperatures predicted by current climate change scenarios will result in an increase in the risk of strike and an elongated blowfly season. However, even for the most rapid warming scenario predictions over the next 70 years, strike is not predicted to occur throughout the winter. Nevertheless, in this latter case, parts of central and southern England are likely to become too hot and dry for strike by L. sericata, to persist in mid-summer. Under these conditions, it is possible that other, more pathogenic Mediterranean agents of myiasis, such as Wolfhartia magnifica, could potentially replace L. sericata. Where the phenology of strike is altered by climate change, as predicted here, significant changes to the timing and frequency of parasite treatments and husbandry practices, such as shearing, will be required to manage the problem. The results suggest that the modelling approach adopted here could be usefully applied to a range of disease systems.     

Interaction of the neuroendocrine system and ovaries in sheep blowflies, Lucilia sericata, sterilized with tepa

ABSTRACT. Female Lucilia sericata Mg. fed a reproductive diet of meat were sterilized by injection of 10μg of tepa. Light microscopy revealed that the medial neurosecretory cells of the brain of sterilized flies were highly active compared with the same cells in untreated flies of similar age and nutritional status. The corpora allata of sterilized flies were also greatly enlarged. The evidence suggests that developing oocytes secrete a substance that inhibits the corpora allata hormone, thereby suppressing brain neurosecretory cell activity and preventing oogenesis in the subsequent oocytes.     

Short-term cold storage of blowfly Lucilia sericata embryos

The developmental rate under low temperatures and cold tolerance were investigated in embryos of the blowfly Lucilia sericata. The larvae of this species are now widely used in maggot debridement therapy. Embryonic development was dependent on temperature, with a lower developmental threshold of 9.0 °C. The duration of the egg stage at a rearing temperature of 25 °C was 14 h, and a low temperature of 12.5 °C successfully prolonged this period to 66 h. Embryonic stages differed markedly in their cold tolerance; young embryos were less tolerant to cold than old ones. Late embryonic stages are suitable for cold storage at 5 °C and the storage for 72 h did not decrease the hatching rate by more than 50%. In the mass‐rearing process required for maggot debridement therapy, either of these two simple protocols would be beneficial.     

Effects of ovarian development and protein deprivation on the activity and locomotor responses of the blowfly, Lucilia sericata, to liver odour

Abstract. The spontaneous activity and locomotor responses to liver odour of females of the blowfly Lucilia sericata Meigen (Diptera: Calliphoridae) were examined in the laboratory using a rocking-box olfactometer. Spontaneous activity levels depended on fly age, previous exposure to protein and recent oviposition history; 1-day-old females, 3-day-old protein-fed females, and females that had recently oviposited exhibited lower levels of spontaneous activity than protein-deprived or fully gravid individuals. Changes in locomotor activity in response to liver odour also depended on previous exposure to protein. In protein-fed flies, activity decreased in response to liver odour and then increased on termination of an odour pulse. This response became more pronounced with age, broadly in line with the expected state of oocyte maturation. This pattern of change was not evident in protein-deprived females. The results are discussed in relation to the relative importance of protein and oviposition-site resources to L. sericata and the resultant age and ovarian development-stage biases evident in field catches.     

Expression of a cGMP compatible Lucilia sericata insect serine proteinase debridement enzyme

Previously, we demonstrated the effectiveness of a research grade recombinant chymotrypsin, derived from the larvae of Lucilia sericata, in "debriding" slough/eschar from venous leg ulcers ex vivo. Furthermore, we were able to formulate this enzyme for successful delivery to in vitro wound healing assays, from a prototype hydrogel wound dressing, and showed that enzyme delivered in this way could degrade wound tissue ex vivo. Recently, to progress biotechnological development of the enzyme as a potential therapeutic product, we explored expression using current good manufacturing practice (cGMP) guidelines, and now report that a recombinant chymotrypsin I zymogen from L. sericata can be expressed in the cGMP acceptable strain of Escherichia coli (BLR-DE3). In addition, the conditions required for purification, refolding and activation of the chymotrypsinogen have been determined. The activated enzyme was stable, and effective in digesting wound slough/eschar tissue. To summarise, we have successfully initiated the production and characterisation of a novel cGMP compatible product for use in future clinical trials.     

Field trial of a compound chromosome strain for genetic control of the sheep blowfly Lucilia cuprina

Summary In 1979–80 a field trial of a compound chromosome (CC) strain of the Australian sheep blowfly Lucilia cuprina was conducted in the isolated Brindabella Valley, N.S.W. New genetic material was introduced into the strain before release by inducing 104 new CC elements by irradiation of recently captured field strains, and combining the resulting strains. Weekly releases, averaging 1.1 million larvae per week, were begun in November 1979 and continued to May 1980. Field-inseminated females were trapped weekly and their genotypes and those of their mates were determined through genetic testing. The proportion of wild X wild matings declined from 16% in December 1979 to 1% in April 1980. During this period the proportion of CC X CC matings rose from 50% to 90%. Larvae sampled from infested sheep had compound chromosomes, indicating that compound chromosomebearing females can successfully oviposit in the field. Trapping of flies resumed at the start of the 1980–81 season, without further releases. Progeny tests revealed the presence of both CC and wild flies. The proportions of CC X CC matings among field-inseminated females were 90% in October, 44% in November, nil in December, and 12% in January. No CC X CC matings were detected in 33 field-inseminated females trapped and tested during April, and 70 tested males reared from myiasis samples in April 1981 proved to be wild type. These results indicate that the CC strain overwintered in the field and strongly suggest that it bred in the field for at least one generation following the spring emergence before being eliminated from the population.     

Reproductive allocation by the blow fly Lucilia sericata in response to protein limitation

Adult females of the anautogenous blow fly Lucilia sericata (Meigen) (Diptera: Calliphoridae) were fed standardized meals of liquidized liver, in quantities shown to range around the minimum required to initiate yolk deposition. Females from each feeding regime were dissected at daily intervals between 4 and 11 days of age; the number and stage of development of all oöcytes were recorded. Once an initial threshold quantity of protein was ingested, yolk deposition was initiated in all available oöcytes. Subsequently, one of two distinct developmental pathways was followed: arrested development in all oöcytes at an early stage of yolk deposition, or more extensive yolk deposition followed by progressive oösorbtion and the maturation of small batches of eggs. The proportion of females showing oösorbtion relative to arrested follicular development increased with increasing protein meal size, suggesting that the difference in response may be triggered by a second protein threshold, either side of which the arrested development or oösorbtion pathways are followed. The behaviour observed may reflect strategies to maximize reproductive output in this short‐lived, resource‐limited insect species. Flies that display arrested development may have sufficient protein to mature few if any complete eggs, but may subsequently be able to mature a full egg‐batch if they obtain further protein meals; this possibility is offset by the risk of death before finding such a meal. Flies that show oöcyte development and oösorbtion produce smaller egg batches more quickly and hence have a higher probability of achieving at least some reproductive output. By initiating yolk deposition in all oöcytes, female L. sericata retain the potential to adopt either developmental pathway, depending on subsequent protein intake.     

A simple, sterile food source for rearing the larvae of Lucilia sericata (Diptera: Calliphoridae)

arvae of the sheep blowfly Lucilia sericata (Meigen) were reared on various mixtures of puréed liver, agar, brewer's yeast and trypsin, in order to develop a simple, sterile, tissue-based diet. Growth and survival rates of larvae reared on a sterile 1:1 mixture of puréed liver with 3% Bacto agar equalled or exceeded those of larvae reared on raw liver. The addition of yeast and/or trypsin to the medium was of no additional benefit. This sterile, homogenous, tissue-based substrate offers a simple, convenient, inexpensive growth medium for rearing larvae for maggot therapy, and for testing the effects of various chemicals and dietary constituents on necrophagous insect larvae. It may be useful, therefore, in studies of myiasis, forensic entomology, and toxicology. This rearing medium also has the advantage that it can be stored for many months at room temperature without progressive decomposition or offensive odour.