Inference of selection pressures that drive insecticide resistance in Anopheles and Culex mosquitoes in Korea.

Pyrethroids are primarily used for mosquito control in Korea. However, high frequencies of mutations conferring resistance to not only pyrethroids but also to other insecticides have been found in mosquito populations. This study aimed to examine the hypothesis that insecticides used outside of public health may play a role in selection. Briefly, the resistance mutation frequencies to three insecticide groups (pyrethroids, organophosphates, and cyclodienes) were estimated in two representative groups of mosquito species (Anopheles Hyrcanus Group and Culex pipiens complex). The relationship between these frequencies and the land-use status of the collection sites was investigated through multiple regression analysis. In the Anopheles Hyrcanus Group, the frequencies of both ace1 (organophosphate resistance) and rdl (cyclodiene resistance) mutations were positively correlated with 'proximity to golf course', possibly be due to the insecticides used for turf maintenance. They also showed positive correlations with field area and rice paddy area, respectively, suggesting the role of agricultural insecticides in the selection of these resistance traits. For the Cx. pipiens complex, the kdr (pyrethroid resistance), ace1, and rdl mutations were positively correlated with the residential area, field, and rice paddy, respectively. Therefore, pyrethroids used for public health could serve as a direct source of resistance selection pressure against kdr, whereas non-public health insecticides may pose primary selection pressure against the ace1 and rdl traits. The current findings suggest that the insecticides used in agriculture and the golf industry play a significant role in mosquito selection, despite variations in the extent of indirect selection pressure according to the mosquito groups and insecticide classes.

Mapping the nutritional landscape in the yellow mealworm: testing the nutrient-mediated life-history trade-offs.

Animals must acquire an ideal amount and balance of macronutrients to optimize their performance, health and fitness. The nutritional landscape provides an integrative framework for analysing how animal phenotypes are associated with multiple nutritional components. Here, we applied this powerful approach to examine how the intake of protein and carbohydrate affects nutrient acquisition and performance in the yellow mealworm (Tenebrio molitor) reared on one of 42 synthetic foods varying in protein and carbohydrate content. Tenebrio molitor larvae increased their food consumption rate in response to nutrient dilution, but this increase was not sufficient to fully compensate for the dilution. Diluting the food nutrient content with cellulose reduced the efficiency of post-ingestive nutrient utilization, further restricting macronutrient acquisition. Tenebrio molitor larvae utilized macronutrients most efficiently at a protein to carbohydrate (P:C) ratio of 1.77:1, but became less efficient at imbalanced P:C ratios. Survivorship was high at high protein intake and fell with decreasing protein intake. Pupal mass and growth rate exhibited a bell-shaped landscape, with the nutritional optima being located around protein-biased P:C ratios of 1.99:1 to 2.03:1 and 1.66:1 to 2.86:1, respectively. The nutritional optimum for development time was also identified at high P:C ratios (1.66:1  to 5.86:1). Unlike these performance traits, lipid content was maximized at carbohydrate-biased P:C ratios of 1:3.88 to 1:3.06. When given a food choice, T. molitor larvae self-composed a slightly carbohydrate-biased P:C ratio of 1:1.24, which lies between the P:C ratios that maximize performance and lipid content. Our findings indicate the occurrence of a nutrient-mediated trade-off between performance and energy storage in this insect.

Behavioural and physiological regulation of protein and carbohydrates in mealworm larvae: A geometric analysis.

Protein-carbohydrate regulation in the larvae of the mealworm beetle (Tenebrio molitor L.) was analyzed using the Geometric Framework for nutrition. In this study, the ingestive and post-ingestive responses were measured from T. molitor larvae that were subjected to choice and no-choice experiments. In the choice experiment, T. moitor larvae were simultaneously presented with one of two protein-biased foods (p35:c7 or p28:c5.6) and one of two carbohydrate-biased foods (p7:c35 or p5.6:c28). T. molitor larvae selected protein and carbohydrate in a ratio close to 1:1 over the first 15 days since the start of the experiment (days 0-15), but exhibited preference for carbohydrate-biased food over the next 15 days. The average protein:carbohydrate ratio selected over days 0-30 was 1:1.24. In the no-choice experiment, T. molitor larvae were restricted to one of seven foods with different protein and carbohydrate content (p0:c42, p7:c35, p14:c28, p21:c21, p28:c14, p35:c7, or p42:c0). On the p0:c42 food, consumption was greatly suppressed and no larvae completed their development. Across a range of these foods except p0:c42, T. molitor larvae consistently over-ate the surplus nutrient in the foods and showed a pattern of nutrient balancing similar to that previously described for other nutritional generalists. Despite having consumed substantially different amounts and ratios of macronutrients as larvae, T. molitor pupae in the no-choice food treatments had similar body nutrient composition, suggesting the presence of strong homeostatic regulation for body nutrient growth. Larval survivorship was significantly lower on two extremely imbalanced foods (p7:c35 and p42:0) than on more balanced foods. T. molitor larvae reared on p7:c35 suffered reduced biomass growth and delayed development compared with those on foods with higher protein content.

Effects of Different Legume Seeds on Individual Performance in the Bean Bug, Riptortus pedestris (Hemiptera: Alydidae).

The bean bug, Riptortus pedestris (Fabricius), is a serious pest of legume crops in East Asia. Here, we report how the consumption of different types of legume seeds affects various aspects of nymphal and adult traits related to fitness in R. pedestris. Two experiments were conducted. In the first experiment, we assessed how R. pedestris nymphs and adults performed on one of 10 different legume seeds: adzuki bean, chickpea, cowpea, two kidney bean varieties (red, white), mung bean, peanut, small black bean, and two soybean varieties (black, yellow). Riptortus pedestris fed on yellow soybean and chickpea performed well in terms of nymphal survivorship and fecundity. However, R. pedestris fed on two kidney bean varieties suffered 100% nymphal mortality and reduced fecundity. Small black bean and black soybean supported high fecundity, but were not suitable for nymphal development. Lipid content was 4- to 11-folds higher in R. pedestris raised on peanuts than those on the others. In the second experiment, we recorded the key parameters of adult performance from R. pedestris assigned to one of nine combinations of three nymphal and three adult diets (yellow soybean, adzuki bean, peanut). Riptortus pedestris raised on yellow soybean during development produced more eggs over their lifetime than those raised on the others. However, this beneficial effect of consuming yellow soybean during development occurred only when yellow soybean was consumed during adulthood. Our data have implications for predicting the occurrence and population dynamics of this pest.

Thermal and nutritional environments during development exert different effects on adult reproductive success in Drosophila melanogaster.

Environments experienced during development have long-lasting consequences for adult performance and fitness. The "environmental matching" hypothesis predicts that individuals perform best when adult and developmental environments match whereas the "silver spoon" hypothesis expects that fitness is higher in individuals developed under favorable environments regardless of adult environments. Temperature and nutrition are the two most influential determinants of environmental quality, but it remains to be elucidated which of these hypotheses better explains the long-term effects of thermal and nutritional histories on adult fitness traits. Here we compared how the temperature and nutrition of larval environment would affect adult survivorship and reproductive success in the fruit fly, Drosophila melanogaster. The aspect of nutrition focused on in this study was the dietary protein-to-carbohydrate (P:C) ratio. The impact of low developmental and adult temperature was to improve adult survivorship. High P:C diet had a negative effect on adult survivorship when ingested during the adult stage, but had a positive effect when ingested during development. No matter whether adult and developmental environments matched or not, females raised in warm and protein-enriched environments produced more eggs than those raised in cool and protein-limiting environments, suggesting the presence of a significant silver spoon effect of larval temperature and nutrition. The effect of larval temperature on adult egg production was weak but persisted across the early adult stage whereas that of larval nutrition was initially strong but diminished rapidly after day 5 posteclosion. Egg production after day 5 was strongly influenced by the P:C ratio of the adult diet, indicating that the diet contributing mainly to reproduction had shifted from larval to adult diet. Our results highlight the importance of thermal and nutritional histories in shaping organismal performance and fitness and also demonstrate how the silver spoon effects of these aspects of environmental histories differ fundamentally in their nature, strength, and persistence.

Combined effects of temperature and macronutrient balance on life-history traits in Drosophila melanogaster: implications for life-history trade-offs and fundamental niche.

Temperature and nutrition are amongst the most influential environmental determinants of Darwinian fitness in ectotherms. Since the ongoing climate warming is known to alter nutritional environments encountered by ectotherms, a precise understanding of the integrated effects of these two factors on ectotherm performance is essential for improving the accuracy of predictions regarding how ectotherms will respond to climate warming. Here we employed response surface methodology to examine how multiple life-history traits were expressed across a grid of environmental conditions representing full combinations of six ambient temperatures (13, 18, 23, 28, 31, 33 °C) and eight dietary protein:carbohydrate ratios (P:C = 1:16, 1:8, 1:4, 1:2, 1:1, 2:1, 4:1, 8:1) in Drosophila melanogaster. Different life-history traits were maximized in different regions in the two-dimensional temperature-nutrient space. The optimal temperature and P:C ratio identified for adult lifespan (13 °C and 1:16) were lower than those for early-life female fecundity (28 °C and 4:1). Similar divergence in thermal and nutritional optima was found between body mass at adult emergence (18 °C and P:C 1:1) and the rate of pre-adult development (28 °C and P:C 4:1). Pre-adult survival was maximized over a broad range of temperature (18-28 °C) and P:C ratio (1:8-8:1). These results indicate that the occurrence of life-history trade-offs is regulated by both temperature and dietary P:C ratio. The estimated measure of fitness was maximized at 23 °C and P:C 2:1. Based on the shape of the response surface constructed for this estimated fitness, we characterized the fundamental thermal and nutritional niche for D. melanogaster with unprecedented detail.

Comparing the impacts of macronutrients on life-history traits in larval and adult Drosophila melanogaster: the use of nutritional geometry and chemically defined diets.

Protein and carbohydrate are the two major macronutrients that exert profound influences over fitness in many organisms, including Drosophila melanogaster. Our understanding of how these macronutrients shape the components of fitness in D. melanogaster has been greatly enhanced by the use of nutritional geometry, but most nutritional geometric analyses on this species have been conducted using semi-synthetic diets that are not chemically well defined. Here, we combined the use of nutritional geometry and chemically defined diets to compare the patterns of larval and adult life-history traits expressed across 34 diets systematically varying in protein:carbohydrate (P:C) ratio and in protein plus carbohydrate (P+C) concentration. The shape of the response surfaces constructed for all larval and adult traits differed significantly from one another, with the nutritional optima being identified at P:C 1:4 for lifespan (P+C 120 g l-1), 1:2 for egg-to-adult viability (120 g l-1), 1:1 for female body mass at adult eclosion (240 g l-1) and lifetime fecundity (360 g l-1), 2:1 for larval developmental rate (60 g l-1) and 8:1 for egg production rate (120 g l-1). Such divergence in nutritional optima among life-history traits indicates that D. melanogaster confined to a single diet cannot maximize the expression of these traits simultaneously and thus may face a life-history trade-off. Our data provide the most comprehensive and nutritionally explicit analysis of the impacts of macronutrients on life-history traits in D. melanogaster and support the emerging notion that the fundamental trade-offs among life-history traits are mediated by macronutrients.

Context-dependent effects of temperature on starvation resistance in Drosophila melanogaster: Mechanisms and ecological implications.

Temperature can modulate the responses of ectotherms to environmental stressors, such as food shortage. Temperature-mediated plasticity in starvation resistance can arise by changes in the amount of energy stored, the speed of energy expenditure, or the threshold energy reserves required for survival. However, few studies have investigated how temperature affects these physiological mechanisms underlying starvation resistance. In this study, we first examined the mechanistic basis of the temperature dependence of starvation resistance in Drosophila melanogaster. We then tested whether the effects of temperature on starvation resistance would depend on diet and developmental stage in this species. We found that exposure to high temperature during starvation significantly reduced the capacity of D. melanogaster to resist starvation. This warming-induced decrease in starvation resistance was mainly caused by faster depletion of body lipids and not by lower threshold lipid content for survival. D. melanogaster exposed to higher temperatures during feeding accumulated more body lipids and thus became more starvation resistant. Such positive effect of high feeding temperature on starvation resistance was pronounced when D. melanogaster consumed carbohydrate-rich diets. D. melanogaster raised as larvae at low temperature (18 °C) had a weaker starvation resistance at adult emergence compared to those raised at higher temperatures (23 and 28 °C). These results demonstrate that the effects of temperature on starvation responses are highly context-dependent in D. melanogaster.

Diet has independent effects on the pace and shape of aging in Drosophila melanogaster.

Studies examining how diet affects mortality risk over age typically characterise mortality using parameters such as aging rates, which condense how much and how quickly the risk of dying changes over time into a single measure. Demographers have suggested that decoupling the tempo and the magnitude of changing mortality risk may facilitate comparative analyses of mortality trajectories, but it is unclear what biologically meaningful information this approach offers. Here, we determine how the amount and ratio of protein and carbohydrate ingested by female Drosophila melanogaster affects how much mortality risk increases over a time-standardised life-course (the shape of aging) and the tempo at which animals live and die (the pace of aging). We find that pace values increased as flies consumed more carbohydrate but declined with increasing protein consumption. Shape values were independent of protein intake but were lowest in flies consuming ~90 µg of carbohydrate daily. As protein intake only affected the pace of aging, varying protein intake rescaled mortality trajectories (i.e. stretched or compressed survival curves), while varying carbohydrate consumption caused deviation from temporal rescaling (i.e. changed the topography of time-standardised survival curves), by affecting pace and shape. Clearly, the pace and shape of aging may vary independently in response to dietary manipulation. This suggests that there is the potential for pace and shape to evolve independently of one another and respond to different physiological processes. Understanding the mechanisms responsible for independent variation in pace and shape, may offer insight into the factors underlying diverse mortality trajectories.

Temperature-driven plasticity in nutrient use and preference in an ectotherm.

Environmental temperature has strong effects on the rate and efficiency of resource use in ectotherms, but little is known about how changes in temperature influence their diet selection patterns. Changes in temperature may alter the balance of nutrients required by ectotherms by affecting metabolism. In response to temperature changes, ectotherms are predicted to express a preference for a specific nutrient (protein or carbohydrate) to match their altered nutrient requirement. Here, we examined the nutritional consequences of mealworm beetles (Tenebrio molitor L.) that were constrained to diets varying in protein:carbohydrate balance (P:C = 1:5, 1:1, or 5:1) or offered a choice between two nutritionally complementary diets (1:5 vs. 5:1) at four different temperatures (20, 25, 30, or 35 °C). Beetles had high mortality and reduced body mass at higher temperatures. Post-ingestive use efficiencies of both protein and carbohydrate decreased as temperature rose. Warming-driven decrease in carbohydrate use efficiencies occurred most profoundly when carbohydrates were consumed excessively. When given a choice, beetles selected protein and carbohydrate equally at 25 and 30 °C, but exhibited a significant preference for carbohydrate at 35 °C. Since carbohydrate is an immediate source of energy, this warming-driven preference for carbohydrate is explicable as an adaptive response of beetles to meet increased energy needs at high temperature. Beetles exposed to 20 °C ate substantially less food, but preferentially consumed carbohydrate over protein possibly to cope with reduced energy intake. The present findings have implications for the impact of temperature on foraging and resource use in ectotherms.

Genetic variation in compensatory feeding for dietary dilution in a generalist caterpillar.

Increasing the rate of food consumption is a common adaptive strategy that allows herbivores to compensate for declines in nutrient concentrations in plant tissues. Herbivores that are better able to compensate for dietary dilution may have selective advantages under nutritionally poor conditions. In order for compensatory feeding to respond to selection, there must be standing heritable variation for this trait. However, empirical data substantiating the adaptive significance and genetic variability of compensatory feeding are rare. By employing a full-sib, split-brood design, this study presents quantitative genetic analyses on the nutrient consumption rates of the generalist caterpillar, Spodoptera exigua, raised on semi-synthetic diets differing in nutrient concentrations. When encountering a diluted diet, caterpillars exhibited a compensatory increase in food consumption rate, but the extent of this increase was not sufficient to fully compensate for dietary dilution. A significant gene-environment interaction for consumption rate indicated that the capacity of caterpillars to compensate for dietary dilution varied across genotypes. The broad-sense heritability of compensatory feeding was 0.51. Caterpillar genotypes with a higher compensatory capacity suffered lower performance losses on the diluted diet than did those with a lower capacity. This study has implications for understanding how herbivores can evolutionarily respond to nutritional challenges.

Balanced intake of protein and carbohydrate maximizes lifetime reproductive success in the mealworm beetle, Tenebrio molitor (Coleoptera: Tenebrionidae).

Recent developments in insect gerontological and nutritional research have suggested that the dietary protein:carbohydrate (P:C) balance is a critical determinant of lifespan and reproduction in many insects. However, most studies investigating this important role of dietary P:C balance have been conducted using dipteran and orthopteran species. In this study, we used the mealworm beetles, Tenebrio molitor L. (Coleoptera: Tenebrionidae), to test the effects of dietary P:C balance on lifespan and reproduction. Regardless of their reproductive status, both male and female beetles had the shortest lifespan at the protein-biased ratio of P:C 5:1. Mean lifespan was the longest at P:C 1:1 for males and at both P:C 1:1 and 1:5 for females. Mating significantly curtailed the lifespan of both males and females, indicating the survival cost of mating. Age-specific egg laying was significantly higher at P:C 1:1 than at the two imbalanced P:C ratios (1:5 or 5:1) at any given age throughout their lives, resulting in the highest lifetime reproductive success at P:C 1:1. When given a choice, beetles actively regulated their intake of protein and carbohydrate to a slightly carbohydrate-biased ratio (P:C 1:1.54-1:1.64 for males and P:C 1:1.3-1:1.36 for females). The self-selected P:C ratio was significantly higher for females than males, reflecting a higher protein requirement for egg production. Collectively, our results add to a growing body of evidence suggesting the key role played by dietary macronutrient balance in shaping lifespan and reproduction in insects.

Macronutrient Balance Modulates the Temperature-Size Rule in an Ectotherm.

Most ectotherms mature at a larger body size in colder conditions, a phenomenon known as the temperature-size rule. While a number of hypotheses have been proposed to explain this rule, little work has been done to understand it from a nutritional perspective. We have used the final-instar caterpillars of Spodoptera litura to investigate how dietary protein∶carbohydrate (P∶C) balance influences the relationship between temperature and body size. The strength and direction of the thermal reaction norm for body size were significantly altered by dietary P∶C balance. The slope of the reaction norm was nearly flat for caterpillars raised on a balanced food ([Formula: see text]) but was significantly negative for those on nutritionally imbalanced foods (1∶5 or 5∶1), especially when carbohydrates were in considerable excess. These nutrient-dependent effects of temperature on body size were caused mainly by corresponding changes in body lipid storage. When allowed to choose between imbalanced diets, caterpillars increased their preference for carbohydrates to meet high energy demands at higher temperatures. The slope of the thermal reaction norm for body size was substantially reduced by such a temperature-driven shift in nutrient preference, indicating that the impact of high temperature on body size was buffered by altered food selection. This study highlights the importance of macronutrient balance as a key factor modulating the relationship between temperature and body size in ectotherms and provides a novel approach for understanding the temperature-size rule.

The genetic basis for mating-induced sex differences in starvation resistance in Drosophila melanogaster.

Multiple genetic and environmental factors interact to influence starvation resistance, which is an important determinant of fitness in many organisms, including Drosophila melanogaster. Recent studies have revealed that mating can alter starvation resistance in female D. melanogaster, but little is known about the behavioral and physiological mechanisms underlying such mating-mediated changes in starvation resistance. In the present study, we first investigated whether the effect of mating on starvation resistance is sex-specific in D. melanogaster. As indicated by a significant sex×mating status interaction, mating increased starvation resistance in females but not in males. In female D. melanogaster, post-mating increase in starvation resistance was mainly attributed to increases in food intake and in the level of lipid storage relative to lean body weight. We then performed quantitative genetic analysis to estimate the proportion of the total phenotypic variance attributable to genetic differences (i.e., heritability) for starvation resistance in mated male and female D. melanogaster. The narrow-sense heritability (h(2)) of starvation resistance was 0.235 and 0.155 for males and females, respectively. Mated females were more resistant to starvation than males in all genotypes, but the degree of such sexual dimorphism varied substantially among genotypes, as indicated by a significant sex×genotype interaction for starvation resistance. Cross-sex genetic correlation was greater than 0 but less than l for starvation resistance, implying that the genetic architecture of this trait was partially shared between the two sexes. For both sexes, starvation resistance was positively correlated with longevity and lipid storage at genetic level. The present study suggests that sex differences in starvation resistance depend on mating status and have a genetic basis in D. melanogaster.

Dietary protein:carbohydrate balance is a critical modulator of lifespan and reproduction in Drosophila melanogaster: a test using a chemically defined diet.

Macronutrient balance is an important determinant of fitness in many animals, including insects. Previous studies have shown that altering the concentrations of yeast and sugar in the semi-synthetic media has a profound impact on lifespan in Drosophila melanogaster, suggesting that dietary protein:carbohydrate (P:C) balance is the main driver of lifespan and ageing processes. However, since yeast is rich in multiple nutrients other than proteins, this lifespan-determining role of dietary P:C balance needs to be further substantiated through trials using a chemically-defined, synthetic diet. In the present study, the effects of dietary P:C balance on lifespan and fecundity were investigated in female D. melanogaster flies fed on one of eight isocaloric synthetic diets differing in P:C ratio (0:1, 1:16, 1:8, 1:4, 1:2, 1:1, 2:1 or 4:1). Lifespan and dietary P:C ratio were related in a convex manner, with lifespan increasing to a peak at the two intermediate P:C ratios (1:2 and 1:4) and falling at the imbalanced ratios (0:1 and 4:1). Ingesting nutritionally imbalanced diets not only caused an earlier onset of senescence but also accelerated the age-dependent increase in mortality. Egg production was suppressed when flies were fed on a protein-deficient food (0:1), but increased with increasing dietary P:C ratio. Long-lived flies at the intermediate P:C ratios (1:2 and 1:4) stored a greater amount of lipids than those short-lived ones at the two imbalanced ratios (0:1 and 4:1). These findings provide a strong support to the notion that adequate dietary P:C balance is crucial for extending lifespan in D. melanogaster and offer new insights into how dietary P:C balance affects lifespan and ageing through its impacts on body composition.

Geometric analysis of nutrient balancing in the mealworm beetle, Tenebrio molitor L. (Coleoptera: Tenebrionidae).

Geometric analysis of the nutritional regulatory responses was performed on an omnivorous mealworm beetle, Tenebrio molitor L. (Coleoptera: Tenebrionidae) to test whether this beetle had the capacity to balance the intake of protein and carbohydrate. We also identified the pattern of ingestive trade-off employed when the insect was forced to balance the costs of over- and under-ingesting macronutrients. When allowed to mix their diet from two nutritionally imbalanced but complementary foods (protein-biased food: p35:c7 or p28:c5.6; carbohydrate-biased food: p7:c35 or p5.6:c28), beetles of both sexes actively regulated their intake of protein and carbohydrate to a ratio of 1:1. When confined to one of seven nutritionally imbalanced foods (p0:c42, p7:c35, p14:c28, p21:c21, p28:c14, p35:c7 or p42:c0), beetles over-ingested the excessive nutrient from these foods to such an extent that all the points of protein-carbohydrate intake aligned linearly in the nutrient space, a pattern that is characteristic of generalist feeders and omnivores. Under the restricted feeding conditions, males ate more nutrients but were less efficient at retaining their body lipids than females. Body lipid content was higher on carbohydrate-rich foods and was positively correlated with starvation resistance. Our results are consistent with the prediction based on the nutritional heterogeneity hypothesis, which links the nutritional regulatory responses of insects to their diet breadth and feeding ecology.

Sex-specific differences in nutrient regulation in a capital breeding caterpillar, Spodoptera litura (Fabricius).

Nutrient requirements by male and female insects are likely to differ, but relatively little is known regarding how sexes differ in their regulation of macronutrient acquisition. The present study reports the results from a laboratory experiment in which behavioural and physiological components of nutrient regulation were compared between male and female caterpillars of Spodoptera litura (Fabricius). When provided with choices between two nutritionally complementary foods (one is a protein-biased diet and the other a carbohydrate-biased diet), both males and females adjusted their food selection to defend an intake target. However, the composition of diet preferred by the two differed, with females selecting significantly more protein than males with no difference in carbohydrate intake between the two. When confined to single diets with varying mixtures of protein and carbohydrate [P:C ratios, expressed as the percentage of diet by dry mass: protein 42%:carbohydrate 0% (p42:c0), p35:c7, p28:c14, p21:c21, p14:c28, p7:c35], females consumed more macronutrients than did males across on all P:C diets except the extremely carbohydrate-biased diet (p7:c35). Under both choice and no-choice feeding condition, such sex differences in nutrient intake were not expressed until late in the feeding stage of the final stadium. Sexes also differed in post-ingestive utilization of ingested nutrients. Females utilized ingested protein for body growth with greater efficiency compared to males, presumably reflecting provisioning their adult needs for protein to develop eggs, whereas males were more efficient at depositing lipids from carbohydrate intake than females.

Can the protein costs of bacterial resistance be offset by altered feeding behaviour?

1. Mounting an immune response is likely to be costly in terms of energy and nutrients, and so it is predicted that dietary intake should change in response to infection to offset these costs. The present study focuses on the interactions between a specialist grass-feeding caterpillar species, the African armyworm Spodoptera exempta, and an opportunist bacterium, Bacillus subtilis. 2. The main aims of the study were (i) to establish the macronutrient costs to the insect host of surviving a systemic bacterial infection, (ii) to determine the relative importance of dietary protein and carbohydrate to immune system functions, and (iii) to determine whether there is an adaptive change in the host's normal feeding behaviour in response to bacterial challenge, such that the nutritional costs of resisting infection are offset. 3. We show that the survival of bacterially infected larvae increased with increasing dietary protein-to-carbohydrate (P:C) ratio, suggesting a protein cost associated with bacterial resistance. As dietary protein levels increased, there was an increase in antibacterial activity, phenoloxidase (PO) activity and protein levels in the haemolymph, providing a potential source for this protein cost. However, there was also evidence for a physiological trade-off between antibacterial activity and phenoloxidase activity, as larvae whose antibacterial activity levels were elevated in response to immune activation had reduced PO activity. 4. When given a choice between two diets varying in their P:C ratios, larvae injected with a sub-lethal dose of bacteria increased their protein intake relative to control larvae whilst maintaining similar carbohydrate intake levels. These results are consistent with the notion that S. exempta larvae alter their feeding behaviour in response to bacterial infection in a manner that is likely to enhance the levels of protein available for producing the immune system components and other factors required to resist bacterial infections ('self-medication').

Lifespan and reproduction in Drosophila: New insights from nutritional geometry.

Modest dietary restriction (DR) prolongs life in a wide range of organisms, spanning single-celled yeast to mammals. Here, we report the use of recent techniques in nutrition research to quantify the detailed relationship between diet, nutrient intake, lifespan, and reproduction in Drosophila melanogaster. Caloric restriction (CR) was not responsible for extending lifespan in our experimental flies. Response surfaces for lifespan and fecundity were maximized at different protein-carbohydrate intakes, with longevity highest at a protein-to-carbohydrate ratio of 1:16 and egg-laying rate maximized at 1:2. Lifetime egg production, the measure closest to fitness, was maximized at an intermediate P:C ratio of 1:4. Flies offered a choice of complementary foods regulated intake to maximize lifetime egg production. The results indicate a role for both direct costs of reproduction and other deleterious consequences of ingesting high levels of protein. We unite a body of apparently conflicting work within a common framework and provide a platform for studying aging in all organisms.

The interactive effects of protein quality and macronutrient imbalance on nutrient balancing in an insect herbivore.

The present study evaluates the interactive effects of protein quality and the ratio of protein to digestible carbohydrate on herbivore performance, various aspects of nutritional homeostasis (feeding and post-ingestive regulation) and food choice. In the first experiment, final-instar caterpillars of generalist-feeding African cotton leafworm Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae) were confined to one of 20 diets varying in protein:carbohydrate (P:C) ratio (35:7, 28:14, 21:21, 14:28 and 7:35) and in the quality of dietary protein (0%, 20%, 50% and 80% replacement of casein-based protein mix with the low-quality plant protein, zein). Results indicated that the negative impact of low-quality protein on survival, development and growth was amplified as the P:C ratio of the diet fell. Consumption differences were the main underlying cause of this response. The effect of low protein quality was to reduce the efficiency of nitrogen utilization and to increase the conversion rate of ingested nutrient to lipid growth, irrespective of P:C ratio. In the second experiment, caterpillars were allowed to self-compose their preferred diet, selecting between one of four high-protein diets (all 35:7, but containing 0%, 20%, 50% or 80% zein) and one of four equal-P:C-ratio diets (21:21, again containing 0%, 20%, 50% or 80% zein). Caterpillars showed a compensatory increase in their self-selected P:C intake in response to the moderate decline in protein quality of the 35:7 food. No such response was demonstrated for the insects presented with 35:7 food with the lowest protein quality. The significance of these findings is discussed within the context of herbivore food selection.