High hemocyte load is associated with increased resistance against parasitoids in Drosophila suzukii, a relative of D. melanogaster

Among the most common parasites of Drosophila in nature are parasitoid wasps, which lay their eggs in fly larvae and pupae. D. melanogaster larvae can mount a cellular immune response against wasp eggs, but female wasps inject venom along with their eggs to block this immune response. Genetic variation in flies for immune resistance against wasps and genetic variation in wasps for virulence against flies largely determines the outcome of any fly-wasp interaction. Interestingly, up to 90% of the variation in fly resistance against wasp parasitism has been linked to a very simple mechanism: flies with increased constitutive blood cell (hemocyte) production are more resistant. However, this relationship has not been tested for Drosophila hosts outside of the melanogaster subgroup, nor has it been tested across a diversity of parasitoid wasp species and strains. We compared hemocyte levels in two fly species from different subgroups, D. melanogaster and D. suzukii, and found that D. suzukii constitutively produces up to five times more hemocytes than D. melanogaster. Using a panel of 24 parasitoid wasp strains representing fifteen species, four families, and multiple virulence strategies, we found that D. suzukii was significantly more resistant to wasp parasitism than D. melanogaster. Thus, our data suggest that the relationship between hemocyte production and wasp resistance is general. However, at least one sympatric wasp species was a highly successful infector of D. suzukii, suggesting specialists can overcome the general resistance afforded to hosts by excessive hemocyte production. Given that D. suzukii is an emerging agricultural pest, identification of the few parasitoid wasps that successfully infect D. suzukii may have value for biocontrol.     

Parasitoid wasp virulence

In nature, larvae of the fruit fly Drosophila melanogaster are commonly infected by parasitoid wasps. Following infection, flies mount an immune response termed cellular encapsulation in which fly immune cells form a multilayered capsule that covers and kills the wasp egg. Parasitoids have thus evolved virulence factors to suppress cellular encapsulation. To uncover the molecular mechanisms underlying the antiwasp response, we and others have begun identifying and functionally characterizing these virulence factors. Our recent work on the Drosophila parasitoid Ganaspis sp.1 has demonstrated that a virulence factor encoding a SERCA-type calcium pump plays an important role in Ganaspis sp.1 virulence. This venom SERCA antagonizes fly immune cell calcium signaling and thereby prevents the activation of the encapsulation response. In this way, the study of wasp virulence factors has revealed a novel aspect of fly immunity, namely a role for calcium signaling in fly immune cell activation, which is conserved with human immunity, again illustrating the marked conservation between fly and mammalian immune responses. Our findings demonstrate that the cellular encapsulation response can serve as a model of immune cell function and can also provide valuable insight into basic cell biological processes.     

An evolutionary conserved role for anaplastic lymphoma kinase in behavioral responses to ethanol

Anaplastic lymphoma kinase (Alk) is a gene expressed in the nervous system that encodes a receptor tyrosine kinase commonly known for its oncogenic function in various human cancers. We have determined that Alk is associated with altered behavioral responses to ethanol in the fruit fly Drosophila melanogaster, in mice, and in humans. Mutant flies containing transposon insertions in dAlk demonstrate increased resistance to the sedating effect of ethanol. Database analyses revealed that Alk expression levels in the brains of recombinant inbred mice are negatively correlated with ethanol-induced ataxia and ethanol consumption. We therefore tested Alk gene knockout mice and found that they sedate longer in response to high doses of ethanol and consume more ethanol than wild-type mice. Finally, sequencing of human ALK led to the discovery of four polymorphisms associated with a low level of response to ethanol, an intermediate phenotype that is predictive of future alcohol use disorders (AUDs). These results suggest that Alk plays an evolutionary conserved role in ethanol-related behaviors. Moreover, ALK may be a novel candidate gene conferring risk for AUDs as well as a potential target for pharmacological intervention.     

Defence strategies against a parasitoid wasp in Drosophila: fight or flight?

Hosts may defend themselves against parasitism through a wide variety of defence mechanisms, but due to finite resources, investment in one defence mechanism may trade-off with investment in another mechanism. We studied resistance strategies against the parasitoid wasp Leptopilina boulardi in two Drosophila species. We found that D. melanogaster had significantly lower physiological resistance against L. boulardi than D. simulans, and hypothesized that D. melanogaster might instead invest more heavily in other forms of defence, such as behavioural defence. We found that when given a choice between clean oviposition sites and sites infested with wasps, both D. melanogaster and D. simulans detected and avoided infested sites, which presumably limits later exposure of their offspring to infection. Unlike D. simulans, however, D. melanogaster laid significantly fewer eggs than controls in the forced presence of wasps. Our findings suggest that D. melanogaster relies more heavily on behavioural avoidance as defence against wasp parasitism than D. simulans, and that this may compensate for a lack of physiological defence.     

ADH enzyme activity and Adh gene expression in Drosophila melanogaster lines differentially selected for increased alcohol tolerance

In Drosophila melanogaster, alcohol dehydrogenase (ADH) activity is essential for ethanol tolerance, but its role may not be restricted to alcohol metabolism alone. Here we describe ADH activity and Adh expression level upon selection for increased alcohol tolerance in different life-stages of D. melanogaster lines with two distinct Adh genotypes: Adh(FF) and Adh(SS). We demonstrate a positive within genotype response for increased alcohol tolerance. Life-stage dependent selection was observed in larvae only. A slight constitutive increase in adult ADH activity for all selection regimes and genotypes was observed, that was not paralleled by Adh expression. Larval Adh expression showed a constitutive increase, that was not reflected in ADH activity. Upon exposure to environmental ethanol, sex, selection regime life stage and genotype appear to have differential effects. Increased ADH activity accompanies increased ethanol tolerance in D. melanogaster but this increase is not paralleled by expression of the Adh gene.     

An ecological interaction between citrus fruit, Penicillium moulds and Drosophila immigrans Sturtevant (Diptera: Drosophilidae)

Abstract. 1. Potential Drosophila breeding sites were collected from a fruit market, and the adults allowed to emerge from them.
2. D.immigrans and D.melanogaster were the species with the highest frequency of emergences from citrus fruit.
3. D.immigrans was especially associated with citrus fruits infected with Penicillium italicum or P.digitatum , two specialist moulds of citrus. D.melanogaster was more often found in uninfected fruit.
4. In the laboratory D.immigrans larvae survived better than D.melanogaster larvae on Penicillium-infected citrus fruit.
5. These adaptations suggest that D.immigrans may have originally evolved as a citrus specialist, becoming a domestic species as these fruits were exploited commercially.     

Behavioural resistance against a protozoan parasite in the monarch butterfly

1. As parasites can dramatically reduce the fitness of their hosts, there should be strong selection for hosts to evolve and maintain defence mechanisms against their parasites. One way in which hosts may protect themselves against parasitism is through altered behaviours, but such defences have been much less studied than other forms of parasite resistance. 2. We studied whether monarch butterflies (Danaus plexippus L.) use altered behaviours to protect themselves and their offspring against the protozoan parasite Ophryocystis elektroscirrha (McLaughlin & Myers (1970), Journal of Protozoology, 17, p. 300). In particular, we studied whether (i) monarch larvae can avoid contact with infectious parasite spores; (ii) infected larvae preferentially consume therapeutic food plants when given a choice or increase the intake of such plants in the absence of choice; and (iii) infected female butterflies preferentially lay their eggs on medicinal plants that make their offspring less sick. 3. We found that monarch larvae were unable to avoid infectious parasite spores. Larvae were also not able to preferentially feed on therapeutic food plants or increase the ingestion of such plants. However, infected female butterflies preferentially laid their eggs on food plants that reduce parasite growth in their offspring. 4. Our results suggest that animals may use altered behaviours as a protection against parasites and that such behaviours may be limited to a single stage in the host-parasite life cycle. Our results also suggest that animals may use altered behaviours to protect their offspring instead of themselves. Thus, our study indicates that an inclusive fitness approach should be adopted to study behavioural defences against parasites.     

Mgat1-dependent N-glycosylation of Membrane Components Primes Drosophila melanogaster Blood Cells for the Cellular Encapsulation Response

In nature, larvae of the fruitfly Drosophila melanogaster are commonly infected by parasitoid wasps, and so have evolved a robust immune response to counter wasp infection. In this response, fly immune cells form a multilayered capsule surrounding the wasp egg, leading to death of the parasite. Many of the molecular mechanisms underlying this encapsulation response are conserved with human immune responses. Our findings suggest that protein N-glycosylation, a common protein post-translational modification of human immune proteins, may be one such conserved mechanism. We found that membrane proteins on Drosophila immune cells are N-glycosylated in a temporally specific manner following wasp infection. Furthermore we have identified mutations in eight genes encoding enzymes of the N-glycosylation pathway that decrease fly resistance to wasp infection. More specifically, loss of protein N-glycosylation in immune cells following wasp infection led to the formation of defective capsules, which disintegrated over time and were thereby unsuccessful at preventing wasp development. Interestingly, we also found that one species of Drosophila parasitoid wasp, Leptopilina victoriae, targets protein N-glycosylation as part of its virulence mechanism, and that overexpression of an N-glycosylation enzyme could confer resistance against this wasp species to otherwise susceptible flies. Taken together, these findings demonstrate that protein N-glycosylation is a key player in Drosophila cellular encapsulation and suggest that this response may provide a novel model to study conserved roles of protein glycosylation in immunity.     

Development of diet-induced insulin resistance in adult Drosophila melanogaster

The fruit fly Drosophila melanogaster is increasingly utilized as an alternative to costly rodent models to study human diseases. Fly models exist for a wide variety of human conditions, such as Alzheimer's and Parkinson's Disease, or cardiac function. Advantages of the fly system are its rapid generation time and its low cost. However, the greatest strength of the fly system are the powerful genetic tools that allow for rapid dissection of molecular disease mechanisms. Here, we describe the diet-dependent development of metabolic phenotypes in adult fruit flies. Depending on the specific type of nutrient, as well as its relative quantity in the diet, flies show weight gain and changes in the levels of storage macromolecules. Furthermore, the activity of insulin-signaling in the major metabolic organ of the fly, the fat body, decreases upon overfeeding. This decrease in insulin-signaling activity in overfed flies is moreover observed when flies are challenged with an acute food stimulus, suggesting that overfeeding leads to insulin resistance. Similar changes were observed in aging flies, with the development of the insulin resistance-like phenotype beginning at early middle ages. Taken together, these data demonstrate that imbalanced diet disrupts metabolic homeostasis in adult D. melanogaster and promotes insulin-resistant phenotypes. Therefore, the fly system may be a useful alternative tool in the investigation of molecular mechanisms of insulin resistance and the development of pharmacologic treatment options.     

Limitations in the use of Drosophila melanogaster as a model host for gram-positive bacterial infection

AIMS: To examine sensitivities of various Drosophila melanogaster strains towards human pathogenic and nonpathogenic gram-positive bacteria. METHODS AND RESULTS: The D. melanogaster Oregon R strain was infected by injecting the thorax with a needle containing Escherichia coli (negative control), Listeria monocytogenes, Staphylococcus aureus (both food-borne pathogens), Listeria innocua, Bacillus subtilis, Carnobacterium maltaromaticum, Lactobacillus plantarum or Pediococcus acidilactici (all nonpathogenic bacteria). Listeria monocytogenes and S. aureus killed the host rapidly compared with the negative control. Infection with L. innocua, B. subtilis or C. maltaromaticum also resulted in a high fly mortality, whereas Lact. plantarum and P. acidilactici resulted in a slightly increased mortality. Four additional D. melanogaster lines, three of which had been selected for heat, cold and desiccation resistance respectively, were subjected to infection by L. monocytogenes, S. aureus and E. coli. Mortality rates were comparable with that of the Oregon R strain. CONCLUSIONS: Use of the injection method shows the limitation of D. melanogaster as a model host for gram-positive bacteria as opportunistic infection by nonpathogenic gram-positive bacteria results in partial or high mortality. In addition, lines of fruit flies resistant to various stress exposures did not show an increased resistance to infection by gram-positive pathogens under the conditions tested. SIGNIFICANCE AND IMPACT OF THE STUDY: This study demonstrates the inadequacy of D. melanogaster infected by the injection method in order to distinguish between virulent and nonvirulent gram-positive bacteria.     

Parasitism affects worker size in the Neotropical swarm-founding social wasp, <Emphasis Type="Italic">Polybia paulista</Emphasis> (Hymenoptera,Vespidae)

Summary We discovered two kinds of parasites, i.e., a strepsipteran, possibly Xenos myrapetrus (Trois) and an undescribed gregarine in the Neotropical swarm-founding paper wasp, Polybia paulista (Ihering). Although proportions of workers that were infected by these parasites varied greatly among colonies analyzed, prevalence of infected workers was recognized. Five external characters were measured and compared among uninfected workers, stylopized (i.e., infected by Strepsiptera) workers and workers that were infected by gregarines. Uninfected workers were significantly larger than stylopized workers, while smaller than workers that were infected by gregarines. Nutrients of stylopized workers may be plundered during their growth period, and consequently their body size may be reduced. However, the gregarines may manipulate host larvae to solicit more food from adults or increase development time of larvae longer, and therefore produce more parasites from a larger host.Received 1 July 2003; revised 1 December 2003; accepted 4 December 2003.     

Do parasitoid preferences for different host species match virulence?

Abstract.  Leptopilina boulardi is a parasitoid wasp specialist of Drosophila larvae of the melanogaster subgroup. In Mediterranean areas, natural populations are highly virulent against their main host Drosophila melanogaster . In Congo, populations are less virulent against D. melanogaster but are able to develop successfully inside the tropical African species Drosophila yakuba . Host preferences are compared between two laboratory isofemale lines of L. boulardi , obtained from populations of Congo and Tunisia, respectively, and differing in virulence levels against D. melanogaster and D. yakuba . Host selection is studied by offering female parasitoids a choice between larvae of the two host species. In agreement with optimal foraging models, the line highly virulent against D. melanogaster shows a clear preference for this host species. The other line, less virulent against D. melanogaster but more virulent against D. yakuba , prefers to oviposit on D. yakuba . Such preferences can be observed after a period of host-patch exploitation only, suggesting that experience plays an important role in the host-selection process. These results evidence the existence of intraspecific variability in preference between two host species in L. boulardi , a major requisite in theoretical models of parasite specialization by the host. They also sustain the hypothesis that intraspecific variation in parasitoid preferences between host species might mirror intraspecific variation in virulence.     

Predation potential of the earwig Euborellia annulipes on fruit fly larvae and trophic interactions with the parasitoid Diachasmimorpha longicaudata

The earwig Euborellia annulipes (Lucas) (Dermaptera: Anisolabididae), a generalist predator, has been observed in fruits infested with fruit fly larvae, which are frequently parasitized by parasitoid wasps. Neither the capacity of earwigs to predate on fruit flies nor intraguild interactions between earwigs and fruit fly parasitoids have been investigated. Here, we studied in laboratory conditions the predation on the fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) by the earwig E. annulipes, and whether parasitism of fruit fly larvae by the parasitoid wasp Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) influences predation by the earwig. We evaluated the predation capacity, functional response and prey preference of E. annulipes for parasitized and non-parasitized fruit fly larvae in choice and no-choice tests. We found that earwigs prey on second- and third-instar larvae and pupae of C. capitata and consumed larger numbers of second-instar larvae, followed by third-instar larvae and pupae. Females prey on larger numbers of fruit flies than did males, regardless of the prey developmental stage, but both sexes exhibited a type II functional response. Interestingly, males killed but did not consume fruit fly larvae more than did females. In no-choice tests, earwig females consumed equal numbers of parasitized and non-parasitized fruit fly larvae. However, in choice tests, the females avoided feeding on parasitized larvae. Subsequent tests with hexane-washed parasitized and non-parasitized larvae showed that putative chemical markings left on fruit flies by parasitoids did not drive the earwig preference towards non-parasitized larvae. These findings suggest that E. annulipes is a potential biological control agent for C. capitata, and that, because the earwig avoids consuming larvae parasitized by D. longicaudata, a combination of the two natural enemies could have an additive effect on pest mortality.     

Opiine parasitoids (Hymenoptera: Braconidae) and biological control of fruit flies (Diptera: Tephritidae) in Australia: Past,present and future

Opiine braconids are parasitoids of the immature stages of frugivorous tephritids. The female wasp lays her eggs into the eggs or larvae of the fruit fly host, where the immature wasp develops before emerging as a next-generation adult from the now dead host pupal case. In support of a new generation of Australian fruit fly parasitoid research, this paper comprehensively reviews what is known about the Australian fruit fly infesting opiines. Based on the most recent taxonomic revision 11 fruit fly infesting opiine species are documented to occur in Australia, but we consider as doubtful the record for Diachasmimorpha longicaudata and consider the record for Fopius illusorius to be tentative without further collections. We identify that the systematics and taxonomy of the Australian native fruit fly infesting opiines are in urgent need of further work. The history of fruit fly biological control in Australia is comprehensively reviewed, including the export of native Australian opiines for fruit fly control elsewhere in the world. Australia was actively involved in three major classical biological control programmes against fruit flies from the turn of the 1900s until the mid-1960s. Despite the introduction of 11 opiine species, plus numerous other natural enemies, only Fopius arisanus established in eastern Australia, where in South-east Queensland it can now cause between 30 and 40% mean parasitism. In addition to the exotic F. arisanus, the native species Diachasmimorpha kraussii and Diachasmimorpha tryoni also cause fruit fly parasitism in agriculturally important crops: both species have also been liberated widely outside of Australia for fruit fly control. Other Australian opiines have not been reared from flies infesting commercial crops and appear biologically restricted to the fruits and environs of Australian east-coast rainforests. The biology literature for D. tryoni and D. kraussii is comprehensively reviewed, while for F. arisanus, already reviewed elsewhere, key literature only is covered. Forward looking, we consider the potential for inoculative or inundative releases of opiines in areas where they do not currently occur to be good, while conservation biological control may help to increase the impacts of parasitoids in areas where they are already established.     

A compound produced by fruigivorous Tephritidae (Diptera) larvae promotes oviposition behavior by the biological control agent Diachasmimorpha longicaudata (Hymenoptera: Braconidae)

Tephritid fruit fly parasitoids use fruit-derived chemical cues and the vibrations that result from larval movements to locate hosts sequestered inside fruit. However, compounds produced by the larvae themselves have not been previously described nor their significance to parasitoid foraging determined. We collected the volatiles from four species of tropical and subtropical Tephritidae: Anastrepha suspensa (Loew), Bactrocera dorsalis Hendel, Bactrocera cucurbitae Coquillett, and Ceratitis capitata (Wiedemann), representing two subfamilies (Dacinae and Trypetinae). Para-ethylacetophenone, an analog of a known tephritid parasitoid attractant, was a major constituent of all four, and was not associated with larvae of another acalypterate fly, Drosophila melanogaster Meigen, or with the calypterate Musca domestica L. It also was present in volatiles from whole, A. suspensa infested fruits of Eugenia uniflora (L.). Para-ethylacetophenone was not necessarily produced as a direct consequence of fruit consumption because it also was detected from larvae that developed in two artificial diets and in spent diets subsequent to larval development. Sensillae on both the antennae and ovipositor of the opiine braconid fruit fly parasitoid, Diachasmimorpha longicaudata (Ashmead) responded to the para-ethylacetophenone in larval volatiles and as a synthetic. Although a potential cue to foraging parasitoids, para-ethylacetophenone showed no long range (>1m) attractiveness to the adult female parasitoid, but did stimulate ovipositor-insertion and oviposition into both a natural (fruit) and an artificial (parafilm) substrate. Thus it may prove useful in colonizing and mass-rearing opine fruit fly parasitoids.     

Evolutionary change in parasitoid resistance under crowded conditions in Drosophila melanogaster

Patterns of investment of limiting resources in such processes as competing for food and defense against natural enemies are shaped by trade-offs and constraints. In Drosophila melanogaster artificial selection for increased resistance to parasitoids results in a correlated decrease in larval competitive ability. Here we ask whether selection for competitive ability leads to a correlated reduction in parasitoid resistance. Replicated lines of D. melanogaster were maintained under crowded or uncrowded conditions for eight generations. As expected, the crowded lines evolved higher competitive ability (when tested against a common strain of fly). But instead of parasitoid resistance decreasing, we found a significant increase, and that this was associated with elevated densities of haemocytes in second-instar larvae. To understand these results we measured a variety of life-history traits in the two sets of lines. We find evidence that directly and indirectly selected changes in competitive ability are due to different mechanisms. We also ask why crowded conditions should select for increased resistance to parasitism, and conclude that it is unlikely to be due to correlated selection for resistance to other natural enemies, but might be due to correlated selection for better wound responses.     

The influence of host fruit morphology on parasitization rates in the Caribbean fruit fly,Anastrepha suspensa

Among the host fruits of the Caribbean fruit fly there are a variety of sizes and shapes. These morphological differences may influence the vulnerability of the larvae to parasites. In the laboratory, Caribbean fruit fly larvae placed in the smaller of 2 different sizes of artificial ‘fruit’ (cloth spheres filled with a diet material) were parasitized at a higher rate by the braconid,Diachasmimorpha longicaudata (Ashmead) when spheres were presented separately. However, when parasites were simultaneously presented with 6 different sizes of ‘fruit’ there was no significant relationship between size and parasitization rate. This may be due to the parasites preference to search for larvae in larger ‘fruit’. In field collections of different species of host fruit, a significant inverse correlation exists between fruit radius and rate of parasitization. However, host fruit size accounts for only about 5% of the variance in yearly parasitization rates.      

Response of Ceratitis capitata, Bactrocera dorsalis, and Bactrocera cucurbitae (Diptera: Tephritidae) to metabolic stress disinfection and disinfestation treatment

Metabolic stress disinfection and disinfestation (MSDD) is a postharvest treatment designed to control pathogens and arthropod pests on commodities that combines short cycles of low pressure/vacuum and high CO2 with ethanol vapor. Experiments were conducted to evaluate the effect of MSDD treatment on various life stages of Ceratitis capitata (Wiedemann), Mediterranean fruit fly; Bactrocera dorsalis Hendel, oriental fruit fly; and Bactrocera cucurbitae Coquillett, melon fly, in petri dishes and in papaya, Carica papaya L., fruit. In some experiments, the ethanol vapor phase was withheld to separate the effects of the physical (low pressure/ambient pressure cycles) and chemical (ethanol vapor plus low pressure) phases of treatment. In the experiments with tephritid fruit fly larvae and adults in petri dishes, mortality was generally high when insects were exposed to ethanol and low when ethanol was withheld during MSDD treatment, suggesting that ethanol vapor is highly lethal but that fruit flies are quite tolerant of short periods of low pressure treatment alone. When papaya fruit infested with fruit fly eggs or larvae were treated by MSDD, they produced fewer pupae than untreated control fruit, but a substantial number of individuals developed nonetheless. This suggests that internally feeding insects in fruit may be partially protected from the toxic effects of the ethanol because the vapor does not easily penetrate the fruit pericarp and pulp. MSDD treatment using the atmospheric conditions tested has limited potential as a disinfestation treatment for internal-feeding quarantine pests such as fruit flies infesting perishable commodities.     

Dolichol levels during development and ageing of Drosophila melanogaster

1. Dolichol levels in the fruit fly, Drosophila melanogaster were determined at the larva and pupa stages and in 1, 10, 20 and 30-day-old flies. 2. Free dolichol increased from 1.2 micrograms/g wet weight in the larvae to 14.9 micrograms/g in 30-day-old flies, while total dolichol increased from 3.4 micrograms/g in the larvae to 21.2 micrograms/g at 30-days-old. 3. Dolichol released after saponification is primarily from dolichyl fatty acid ester, which accounts for up to 65% of the total dolichol. 4. The major dolichol homologs, which remain relatively constant throughout development and ageing in D. melanogaster, are C-80, C-85 and C-90, which represent approximately 7%, 60% and 33%, respectively.     

Drosophila melanogaster larvae make nutritional choices that minimize developmental time

Organisms from slime moulds to humans carefully regulate their macronutrient intake to optimize a wide range of life history characters including survival, stress resistance, and reproductive success. However, life history characters often differ in their response to nutrition, forcing organisms to make foraging decisions while balancing the trade-offs between these effects. To date, we have a limited understanding of how the nutritional environment shapes the relationship between life history characters and foraging decisions. To gain insight into the problem, we used a geometric framework for nutrition to assess how the protein and carbohydrate content of the larval diet affected key life history traits in the fruit fly, Drosophila melanogaster. In no-choice assays, survival from egg to pupae, female and male body size, and ovariole number – a proxy for female fecundity – were maximized at the highest protein to carbohydrate (P:C) ratio (1.5:1). In contrast, development time was minimized at intermediate P:C ratios, around 1:2. Next, we subjected larvae to two-choice tests to determine how they regulated their protein and carbohydrate intake in relation to these life history traits. Our results show that larvae targeted their consumption to P:C ratios that minimized development time. Finally, we examined whether adult females also chose to lay their eggs in the P:C ratios that minimized developmental time. Using a three-choice assay, we found that adult females preferentially laid their eggs in food P:C ratios that were suboptimal for all larval life history traits. Our results demonstrate that D. melanogaster larvae make foraging decisions that trade-off developmental time with body size, ovariole number, and survival. In addition, adult females make oviposition decisions that do not appear to benefit the larvae. We propose that these decisions may reflect the living nature of the larval nutritional environment in rotting fruit. These studies illustrate the interaction between the nutritional environment, life history traits, and foraging choices in D. melanogaster, and lend insight into the ecology of their foraging decisions.