Food consumption and utilization responses to dietary dilution with cellulose and water by velvetbean caterpillars, Anticarsia gemmatalis

Abstract Dilution of an artificial diet with water or cellulose to nutrient levels of 32% (undiluted), 19% and 10% fresh weight (fw) resulted in increased fw and dry weight (dw) food consumption (both absolute amounts and weight-relative rates) by velvetbean caterpillars, Anticarsia gemmatalis Hübner. Despite these increases, the absolute amount and relative rate of nutrient intake by the caterpillars declined with dilution, as did their dw gain and dw (% fw) and lipid (% dw) contents. The proportion of consumed food (dw, including cellulose) that was digested and absorbed declined with increased dietary cellulose; however, the proportion of consumed nutrients (dw, excluding cellulose) that was digested and absorbed was not affected by the presence of cellulose, although it increased slightly (but significantly) with dietary water. The efficiency with which the absorbed nutrients were converted to biomass energy showed a negative relationship with the various measures of food consumption and thus a positive relationship with dietary nutrient level.
The compensatory increases in feeding mitigated the deleterious impact of dietary dilution on growth, which would have declined further without the increased feeding. These data suggest that food consumption is regulated primarily through an evaluation of food nutrient level, and that the metabolic costs associated with the increased consumption and processing of food may be substantial, especially on the most diluted diets. Another hypothesized cost of increased feeding, reduced efficiency of digestion and absorption as food presumably passes more rapidly through the gut, was not detected.     

Effects of potassium levels over the range occurring in host plants on the growth of an insect herbivore (Manduca sexta)

Third instar tobacco hornworms (Manduca sexta L.: Sphingidae) on low dietary potassium had a lower relative growth rate than individuals on diets with potassium concentrations reflecting those in host-plants, due to decreased consumption rate, lower efficiencies of conversion of ingested and digested food (ECI and ECD), and a prolonged growth/feeding phase. Furthermore, these larvae, when placed on a diet with a moderate potassium concentration through the fourth stadium, ended up being smaller due to lower ECI and less biomass gained, and had a prolonged growth phase, which suggest an irreversible cost of the previous low potassium diet. Third instar hornworms on high potassium diets had lower ECI and ECD, and they had a prolonged growth phase. These individuals, when placed on a moderate potassium diet in the fourth stadium, gained less biomass, than those previously offered hostplant-like-potassium diets. Body potassium concentrations (% dw) at the end of the third stadium were similar among treatment groups. With increasing potassium concentrations in the diet, utilization efficiencies of potassium decreased and potassium concentrations in the frass increased. Correspondingly, water content (% fw) of the newly-molted fourth instar larvae declined with increasing potassium, indicating a passive loss of water during potassium excretion. Low and high dietary potassium reduced survivorship of third instar larvae; fourth instar caterpillars previously fed the low potassium diet also had poor survivorship. We conclude that, within the normal range of potassium concentrations in the hostplants, caterpillar performance is largely unaffected by potassium concentration, but that potassium-poor and potassium-rich diets, such as those hornworms may sometimes experience, can reduce growth and survivorship.     

Effects of dextrin and cellulose on feeding and reproduction in female brown-banded cockroaches, Supella longipalpa

Abstract Food consumption and reproduction were compared through two gonotrophic cycles in female brown-banded cockroaches, Supella longipalpa (F.), fed a standard rat food diet which had been serially diluted with methyl cellulose, alpha cellulose or dextrin. Females did not increase daily intake to compensate for dilution, and reproductive rate was highly dependent upon the degree of dilution of the diet. With increasing dextrin concentrations, digestibility increased, but reproductive rate, size and mass of oothecae, and efficiency of food utilization progressively decreased.
On diets containing 75% rat food and 25% methyl cellulose, females consumed minimal amounts of food and 80% of the females did not mate or produce oothecae within 30 days. On identical diets containing 25% alpha cellulose, food digestibility decreased and females ate significantly more than females fed 25% methyl cellulose, but less than females fed rat food. Many (65%) females fed 25% alpha cellulose produced oothecae which were smaller and took longer to form than in control females fed on rat food. At concentrations higher than 50% of either methyl or alpha cellulose daily intake was further reduced and females died rapidly.     

Compensatory feeding in response to variable food quality by Melanoplus differentialis

Abstract. The behavioural mechanisms driving compensatory food consumption in response to dietary dilution as well as the relationship between feeding time and food residence time (i.e. digesta retention time in the gut) were studied using the non-diapausing strain of the grasshopper Melanoplus differentialis (Thomas) (Orthoptera: Acrididae). 3-day-old, sixth-instar nymphs and 3-day-old adults were fed artificial diets containing 1%, 3% and 5% total nitrogen (N) at 30C, LD 14:10h; the feeding behaviour was recorded using electronic monitoring devices connected to microcomputers for 24 h. The percentage of time spent feeding increased linearly as diets were diluted using non-digestible cellulose from 5% to 1% N. This response was due to an increase in the number of meals while the meal duration of a feeding bout was unaltered. Sixth-instar nymphs spent about 40% more time feeding than the larger adults. The increased feeding time in nymphs resulted from both more frequent feeding bouts and longer meal duration. Feeding time and food residence time were highly negatively related.     

Do food protein and carbohydrate content influence the pattern of feeding and the tendency to explore of forest tent caterpillars?

This study examines whether the ratio of protein to carbohydrate affects the timing of meals and the propensity to explore of forest tent caterpillars (Malacosoma disstria). The behavior of fourth instar caterpillars was observed on three semi-defined artificial diets varying in protein (p)-carbohydrate (c) ratio. These diets were (a) p14:c28, (b) p28:c14, and (c) p35:c7. The probability of initiating feeding at first contact with the food and the duration of the first feeding event did not vary across diets, suggesting not much difference in phagostimulatory power. There was also no difference in the total time spent eating, at rest and in motion between diets. However, the timing and duration of meals varied significantly; more short meals were observed on the carbohydrate-biased diet. The duration of pauses between meals also increased with food protein content. Furthermore, caterpillars on the carbohydrate-biased diet were more likely to leave the trail leading to the known food source and to discover a second food source, suggesting that protein deprivation promotes exploration. These findings shed insight into the physiological responses to protein and carbohydrate ingestion and demonstrate how post-ingestive effects can favor consumption of foods containing protein without invoking an explicit mechanism of independent nutrient regulation, but simply by influencing the pattern of feeding and the propensity to explore.     

Food and water economy and its relation to growth in fifth-instar larvae of the tobacco hornworm, Manduca sexta

Fifth-instar larvae of Manduca sexta were reared on artificial diet at 25°C. Daily measurement of the wet and dry weights of insects, food eaten and faeces produced, allowed the construction of detailed budgets for food and water economy.Approximate digestibility (AD) was about 60% both in dry weight and energetic terms. This did not change significantly on successive days of the instar, despite the very considerable changes that occur in the size of the insect and the amount of food eaten during the same period. Two indices of conversion efficiency (ECI and ECD) also showed only insignificant changes during this time. ECI and ECD were both very high: at their maximal values, ECI was 43% (51% in energy terms) and ECD was 72% (81% in energy terms).Correction of the nutritional indices for the presence of food in the gut reduced their value slightly in all cases.Most of the water requirements of the larva were provided by the diet. The estimated gain from metabolic water was between 3.5 and 10% of the total water input. Faeces accounted for most of the water lost from the body; under our conditions transpirational losses were very small. Almost half of the water taken up with the food was retained in the body.The gut contents are more hydrated than the food. The faeces contain less water than the gut contents. It is suggested that the caterpillars recycle water by absorbing it from the faeces and returning it to the contents of the midgut. A quantitative model of such water movement within the body is presented. The data provide indirect evidence that caterpillars may optimise the retention time of food in the gut (and thus the AD) in order to maximise the absolute rate of nutrient uptake from it.     

Compensatory responses of the fall armyworm (Spodoptera frugiperda) when fed water- and cellulose-diluted diets

Abstract Caterpillars of the noctuid moth, Spodoptera frugiperda (J. E. Smith), reared on artificial diets diluted with cellulose and water, increased fresh weight (fw) consumption 2.2–2.5-fold over those on the undiluted diet. At moderate levels of water- or cellulose-dilution, the increased consumption, combined with increased digestion and absorption of nutrients (ADNU), sufficiently compensated for the reduced nutrient intake to achieve pupal biomass equivalent to that of pupae from the undiluted diet. At higher levels of water- and cellulose-dilution, fw consumption and ADNU increased further but pupal dry weight declined on the water-diluted diets. At each level of dilution fw consumption and ADNU increased similarly on the water- and cellulose-diluted diets, but biomass gain was reduced on the water- compared with the cellulose-diluted diets. This was due in part to lowered food conversion efficiency on the water-diluted diets, possibly caused by increased costs of metabolizing the wetter diets. Our data support the hypothesis that consumption rates are regulated by a nutrient feedback mechanism. Our data also suggest that digestive enzyme activity is correlated with consumption. Furthermore, the cost of increased consumption rates on diets of reduced energetic value may constitute a substantially greater energy expenditure than previously believed. However, this cost was insufficient to reduce relative growth rates but was apparently manifested in lowered lipid accumulation.     

Combined effects of allelochemical-fed and scarce prey on the generalist insect predator Podisusmaculiventris

1. The simultaneous effects of allelochemicals ingested by herbivorous insect prey and prey scarcity on the performance of a generalist insect predator were examined.
2. Fifth-instar predatory stinkbugs ( Podisus maculiventris : Pentatomidae) were fed caterpillars ( Manduca sexta : Sphingidae) in three prey scarcity treatments: every day (unlimited amount), one caterpillar every third day, one caterpillar every fifth day. The caterpillars were fed either a plain diet or a diet containing rutin, chlorogenic acid and tomatine, which are three of the major allelochemicals in tomato leaves ( Lycopersicon esculentum : Solanaceae), the preferred food of these caterpillars.
3. Food consumed, efficiency of conversion of ingested food to biomass (ECI), biomass gained, stadium duration and relative growth rate (RGR) of predators were negatively affected by prey scarcity. The allelochemicals negatively affected food consumed and ECI.
4. There were prey scarcity by allelochemical interactions for ECI, biomass gained and RGR. For ECI, the allelochemicals had a greater negative impact on the predatory stinkbugs when prey were scarce. When prey diet contained allelochemicals, biomass gained and RGR declined more steeply with increased prey scarcity. There was an allelochemical by predator gender interaction for biomass gained. Allelochemicals had no effect on biomass gained by female stinkbugs, whereas biomass gained declined more steeply with increased prey scarcity for male stinkbugs fed caterpillars containing allelochemicals than for males fed control caterpillars.     

Three hundred and fifty generations of extreme food specialisation: testing predictions of nutritional ecology

We used a strain of diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae), that had been reared for approximately 350 generations in a precisely characterised environment to test hypotheses regarding the influence of nutritional heterogeneity on the evolution of nutrient regulatory responses. Caterpillars were maintained with ad libitum access to a diet that emulated that of an extreme nutritional specialist, comprising a homogeneous food of fixed nutrient composition. We measured performance (survival, development rate, and pupal mass), as well as the protein and carbohydrate intake of individual caterpillars confined to one of a range of single foods differing in their protein, carbohydrate, and water content. In a separate experiment, we measured the amount and balance of protein and carbohydrate self-selected by caterpillars presented with nutritionally complementary foods. Results showed a close fit with three of four predictions about the nutritional responses of 'nutrient specialist' feeders: (1) survival, development rate, and pupal mass were highest for animals given diets with the protein:carbohydrate composition of the ancestral culture diet, and dropped off sharply with higher and lower protein:carbohydrate balance, (2) caterpillars coped poorly with dietary dilution by water, irrespective of the macronutrient balance, and (3) the self-selected intake point corresponded with the macronutrient balance that gave peak performance (i.e., that of the ancestral culture diet). The fourth prediction, that caterpillars would be disinclined to over-ingest nutrients on imbalanced diets, was at best weakly met. We hypothesise that the evolution and maintenance of the specialist strategy might, paradoxically, require some degree of environmental heterogeneity.     

The interplay between nutrient balancing and toxin dilution in foraging by a generalist insect herbivore

Food mixing by herbivores is thought to balance nutrient intake and possibly dilute secondary metabolites characteristic of different host plant species. Most empirical work on insect herbivores has focused on nutrient balancing in laboratory settings. In this study, we characterize food mixing behaviour of the caterpillar Grammia geneura (Strecker) (Lepidoptera: Arctiidae) in nature and use the observed patterns to design ecologically relevant experiments that reveal the relative importance of these processes in food-switching behaviour. Our design involved both choice and no-choice experiments with chemically defined diets in which primary nutrients and secondary metabolites were manipulated in tandem. We analysed two stages in the process of food-switching behaviour: leaving food and accepting new food. In nature, an individual's rate of leaving host plants was positively associated with its probability of rejecting plant species most recently eaten, but not related to its probability of accepting different host plant species. Furthermore, an individual's leaving rate was negatively related to its average feeding bout duration. This relationship resulted partly from variation in the response of individuals to nutrient imbalance and partly from shortened feeding bouts prior to switching, suggesting that a decline in feeding excitation preceded searching for food that differed from that most recently eaten. Laboratory experiments with synthetic diets indicated the importance of secondary metabolites in the decline in feeding excitation prior to switching. Preference for new food depended strongly on secondary metabolites in a manner consistent with toxin dilution. This is the first experimental evidence for the process of toxin dilution in caterpillars, and for the combined influence of nutrients and secondary metabolites on their foraging patterns in nature. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Caterpillars' compensatory feeding response to diluted nutrients leads to toxic allelochemical dose

Many herbivores increase their consumption rate as dietary nutrient concentration declines. This compensatory response can mitigate the fitness-lowering impact of reduced food quality, but little is known about its costs. In this study we tested the hypothesis that one cost to a faster consumption rate can be the ingestion of a toxic dose of an allelochemical occurring in the food. We fed velvetbean caterpillars a diet with progressively diluted nutrient levels but containing the same concentration (% fresh mass, fm) of caffeine, a methylxanthine alkaloid. Larvae compensated for the reduced nutrient level, with those fed the most diluted diet increasing their biomass-relative consumption rate (fm) 2.6-fold over larvae fed the undiluted diet. Consequently, their rate of caffeine ingestion increased to a pharmacologically effective dose, interfering with food utilization, slowing growth, reducing subsequent feeding and lowering survival. These results suggest that greater allelochemical ingestion can be one cost of an increased consumption rate, although additional studies with other allelochemicals and species are necessary to more broadly evaluate whether insects can adaptively balance their intake of nutrients and allelochemicals through adjustments in consumption rate. In addition, these results highlight the importance of measuring consumption rates of allelochemicals and other ingested biocides, not just their dietary concentration, when assessing efficacy against herbivores.     

Influence of dietary nutritional composition on caterpillar salivary enzyme activity

Caterpillars are faced with nutritional challenges when feeding on plants. In addition to harmful secondary metabolites and protein- and water-limitations, tissues may be carbohydrate-rich which may attenuate optimal caterpillar performance. Therefore, caterpillars have multiple strategies to cope with surplus carbohydrates. In this study, we raise the possibility of a pre-ingestive mechanism to metabolically deal with excess dietary sugars. Many Noctuid caterpillars secrete the labial salivary enzyme glucose oxidase (GOX), which oxidizes glucose to hydrogen peroxide and gluconate, a nutritionally unavailable carbohydrate to the insect. Beet armyworm, Spodoptera exigua, larvae were restricted to diets varying in protein to digestible carbohydrate (P:C) ratio (42p:21c; 33p:30c; 21p:42c) and total nutrient concentration (42% and 63%). High mortality and longer developmental time were observed when caterpillars were reared on the C-biased, P-poor diet (21p:42c). As the carbohydrate content of the diet increased, caterpillars egested excess glucose and a diet-dependent difference in assimilated carbohydrates and pupal biomass was not observed, even though caterpillars restricted to the C-biased diet (21p:42c) accumulated greater pupal lipid reserves. Larval labial salivary GOX activity was also diet-dependent and gluconate, the product of GOX activity, was detected in the frass. Unexpectedly, GOX activity was strongly and positively correlated with dietary protein content.     

The effects of increased dietary cellulose on the anatomy, physiology and behaviour of captive water voles, Arvicola terrestris (L.) (Rodentia: Microtinae)

1. Wild water voles and captives on high fibre diets had significantly longer and heavier digestive tracts (especially the caecum) than captives fed a low fibre diet. 2. Increasing dietary cellulose increased food intake, faecal output and metabolic rate, and decreased digestibility. Some but not all added cellulose was digested. 3. The passage of food through the digestive tract was significantly faster on the diet with added cellulose. 4. Coprophagy included eating directly from the anus and eating faecal pellets from the floor of the cage. The frequency of the latter type increased significantly for animals on a high fibre diet.     

Compensatory feeding response of the slug Sarasinula plebeia to dietary dilution

Summary Like many polyphagous herbivores, individuals of Sarasinula plebeia (Fischer) (Soleolifera: Veronicellidae) consume a variety of plant species that may differ in nutritional content. In this study we determined the ability of these slugs to compensate for such variation in diet composition. Dilution with water of an agar-based diet containing commercial guinea pig food or carrot root to obtain dry weights (dw) of 90, 70, 40 and 10% of diet fresh weight (fw), caused immature slugs to consume increasingly more fresh weight of food [as much as 4.7-(guinea pig) to 6.1-fold (carrot) more]. Dry weight consumption and body mass-relative dry weight consumption rate also increased at intermediate dilutions, buth with further dilution, dry weight intake declined despite the greater fresh weight consumption. At each dilution level, slugs fed the guinea pig diet consumed from ca. 5-to 6.4-fold more fresh weight than the carrotfed slugs. The former grew substantially, with their final biomass and body mass-relative growth rate varying curvilinearly with diet % dw. If these slugs had not fed more but instead maintained the same fresh weight consumption as slugs in the 90% dw tretments, without altering food utilization efficiencies, then their biomass gain in the 70, 40 and 10% dw treatments would have been only about 62, 43, and 21%, respectively, of the values actually attained. In contrast, carrot-fed slugs did not grow and were only able to maintain their initial biomass. For each diet, slug tissue water (% fw) was highest in the most diluted treatment but did not differ significantly among the other dilution levels. Approximate digestibility of the carrot diet was highest at intermediate dilution levels (ca. 75% of ingested food was digested and absorbed); for the guinea pig diet, this efficiency declined linearly from about 66% to 59% with increased dilution. For slugs that grew (i.e., those fed the guinea pig diet), effeciences of converting digested (29–52%) and ingested (18–33%) food to dry biomass were both curvilinearly related to diet % dw. Thus, S. plebeia, like many other herbivores, has the capacity to increase food consumption substantially inresponse to reduced dietary nutrient level, allowing the slugs to cope with variable nutrient content in their food plants.     

Dietary protein supplementation and its consequences for intake,digestion, and physical activity of a carnivorous marsupial,Sminthopsis crassicaudata

Diet regulation behavior can mediate the consequences of imbalanced diets for animal well‐being, particularly for captive species that have little dietary choice. Dasyurids (carnivorous marsupials) are of conservation concern in Australia, and many species are in captive breeding programmes. However, their nutrient targets and dietary regulation behaviors are poorly understood, a limitation that may decrease the breeding success and well‐being of captive animals. We tested how dietary protein content influenced the intake and utilization of nutrients, physical activity, and body mass of fat‐tailed dunnarts Sminthopsis crassicaudata. Twelve adult dunnarts from six sibling pairs (one female and one male per pair) were provided ad libitum access to three diets in a repeated measures design: cat food, cat food supplemented with raw lean beef (1:1), and cat food supplemented with cooked lean beef (1:1). Food intake, activity level, and fecal output were measured daily. Dunnarts significantly decreased food intake, increased protein digestion, and physical activity, but body mass was unchanged when on the high‐protein diet compared to the normal cat food diet. These observations suggest a capacity of dunnarts to maintain constant body mass using a dynamic balance of feeding, digestion, and activity. We also found a significant effect of family, with differences between families as large as the difference between the diet treatments, suggesting a genetic component to diet selection. The nutrient regulation responses of dunnarts to high‐protein diets and the strong family effects provide important messages for the management of populations of small carnivores, including the aspects of dietary manipulation and conservation of genetic diversity.     

Clonidine increases food and protein consumption in rats

Rats were given clonidine or its diluent, and allowed to eat freely from two isocaloric diets that differred in protein or carbohydrate content. Low clonidine doses (25–50 μg/kg) significantly increased total food and protein intake by rats given access to a high and a low protein diet. Several pairs of diets, differing in protein contents, were tested; clonidine's effect was greatest when a diet containing 30–45% protein was paired with one that was very low (5%) in protein. Higher clonidine doses (200 μg/kg) failed to modify either total food or protein intake. Clonidine had no effects on food or nutrient intake among animals given access to diets that differed in carbohydrate content (25 or 70% carbohydrate, plus 25% protein). In rats given access to only one diet, clonidine administration decreased food consumption when the diet was low in protein (5%), but increased consumption when the diet contained 25 or 50% protein. These data suggest that central noradrenergic synapses participate in the mechanisms controlling appetites for proteins. Clonidine may enhance protein intake by stimulating presynaptic alpha receptors, thus diminishing central noradrenergic tone. This effect on noradrenergic transmission is probably partly overcome by protein consumption, which increases brain tyrosine levels and thus can accelerate norepinephrine synthesis. Clonidine or related drugs may be useful clinically in treating diseases characterized by impaired appetite or increased need for protein.     

Consumption of a toxic food by caterpillars increases with dietary exposure: support for a role of induced detoxification enzymes

We examined whether there is a link between feeding behavior and detoxification enzyme activity, which increases over time in proportion to the ingested dose of toxic allelochemicals. We hypothesized that, for insects consuming a toxic but nondeterrent food, consumption would initially decline but then increase after continuous exposure in conjunction with the induction of detoxification enzyme activity. We tested this hypothesis by observing the feeding behavior of fall armyworms (Spodoptera frugiperda) when offered diets containing indole 3-carbinol (I3C), a well-known inducer of detoxification enzymes in this species. The I3C concentrations used were toxic but nondeterrent, thereby avoiding any confounding influence of preingestive responses. The feeding pattern of caterpillars previously unexposed to dietary I3C (i.e. with basal detoxification enzyme activity) was substantially altered when offered I3C-containing diets. Compared with control caterpillars fed I3C-free diets, they exhibited fewer feeding bouts and had a greater proportion of long (> 60 min) non-feeding pauses between bouts. In contrast, the feeding pattern of caterpillars pre-exposed to an I3C diet (i.e. with induced detoxification enzyme activity) did not differ from that of control caterpillars. We conclude that there is a relationship between feeding behavior and detoxification activity in these caterpillars, and propose two postingestive mechanisms that could link these processes.     

Water ingestion by rats fed a high-salt diet may be mediated, in part, by visceral osmoreceptors

After surgical removal of all salivary secretions ("desalivation"), rats increase their consumption of water while eating dry laboratory chow. In the present experiments, desalivated rats drank even more water while they ate "powdered" high-salt food (i.e., <15-mg food particles). The Na+ concentration of systemic plasma in these animals was not elevated during or immediately after the meal, which suggests that cerebral osmoreceptors were not involved in mediating the increased water intake. A presystemic osmoregulatory signal likely stimulated thirst because the Na+ and water contents of the gastric chyme computed to a solution approximately 150 mM NaCl. In contrast, desalivated rats drank much smaller volumes of water while eating "pulverized" high-salt food (i.e., 60-140-mg food particles), and the fluid mixture in the gastric chyme computed to approximately 280 mM NaCl solution. These and other findings suggest that the NaCl ingested in the powdered high-salt diet was dissolved in the gastric fluid and that duodenal osmoreceptors (or Na+-receptors) detected when the concentration of fluid leaving the stomach was elevated after each feeding bout, and promptly stimulated thirst, whereupon rats drank water until the gastric fluid was diluted back to isotonicity. However, when rats ate the pulverized high-salt diet, much of the NaCl ingested may have been embedded in the gastric chyme and therefore was not accessible to visceral osmoreceptors once it emptied from the stomach. Consistent with that hypothesis, fluid intakes were increased considerably when desalivated rats drank 0.10 M NaCl instead of water while eating either powdered or pulverized high-salt food.     

Nutrient utilization by caterpillars of the generalist beet armyworm, Spodoptera exigua

Abstract.  Beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), caterpillars are destructive crop pests responsible for considerable annual economic losses. These generalist herbivores are challenged with a diversity of dietary choices that can impact their survival, development and fecundity. In the present study, dietary choices of S. exigua caterpillars, based on the protein to digestible carbohydrate (P : C) ratio of the food, and the impact of nutritionally unbalanced foods on caterpillar performance are assessed. In choice experiments, individual third-instar caterpillars are offered simultaneously a P-biased and a C-biased food until pupation. Caterpillars feed nonrandomly and select a slightly P-biased diet (22P : 20C). In no-choice experiments, second instar caterpillars are reared until pupation on diets ranging in P : C ratio from extremely P- to extremely C-biased. High mortality and delayed development are observed on the C-rich, P-poor diets, highlighting the potential deleterious effects of excess carbohydrates and the importance of protein for growth and development. Diet-dependent differences in pupal weight or pupal lipid reserves are not observed. This contrasts with closely-related Spodoptera species where pupal mass and lipid stores increase on C-rich, P-poor diets. On the extremely P-biased diet, performance is similar to that of individuals reared on the self-selected diet, suggesting that these caterpillars may efficiently be deaminating excess amino acids to generate carbon skeletons, which are shunted into lipid biosynthesis. Spodoptera exigua caterpillars exhibit flexible and efficient pre-ingestive nutrient intake regulation and post-ingestive utilization, allowing these generalist feeders to cope with the heterogeneous diets they may encounter.     

Compensatory increases in food consumption and utilization efficiencies by velvetbean caterpillars mitigate impact of diluted diets on growth

Fresh weight (fw) food consumption by caterpillars of Anticarsia gemmatalis Hübner (Lepidoptera: Noctuidae) increased almost 2-fold as the nutrients in an artificial diet were increasingly diluted with water (diets contained 65, 79, 86 or 89% fw water). Nonetheless, dry weight (dw) relative consumption rate (RCR) declined with diet dilution. The efficiency at which the consumed food is digested and assimilated (approximate digestibility, AD) increased on the 3 diluted diets, and the efficiency at which digested food is converted to biomass (ECD) increased on the 79 and 86% fw diets. As a consequence, dw gained and relative growth rate (RGR), which is the product of RCR x AD x ECD, on the 79% fw diet were similar to those on the 65% fw diet, but they declined on the more diluted diets. Relative nitrogen consumption rate also declined with dietary dilution, but this was compensated by an increase in nitrogen utilization efficiency such that the product of these, relative nitrogen accumulation rate, was similar on all four diets. Insect lipid content declined from 32% on the undiluted diet to 13% dw on the most diluted diet, and was primarily responsible for the decline in RGR. The increases in fw consumption and AD, while not preventing a decline in RGR on the two most diluted diets, mitigated the impact of dietary dilution (e.g., without these increases, RGR on the most diluted diet would have been only 43% of that attained). These results indicate that the consumption and utilization of food are dynamic processes, and that caterpillars of A. gemmatalis, like many other insects, exhibit compensatory responses to changes in dietary quality.

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Résumé Peu de travaux ont examiné l'aptitude des insectes à modifier leur consommation et l'utilisation des aliments en fonction des variations qualitatives et quantitatives connues de l'aliment. On a rarement examiné si ces modifications sont adaptatives, si elles maintiennent le taux de croissance et le niveau des ressources, ou si elles limitent au moins les effets nuisibles du changement d'aliment. En utilisant une technique gravimétrique, nous avons mesuré le gain des poids, l'aliment consommé et les excréments produits par des chenilles de A. gemmatalis Hübn. (Lep, Noctuidea), important ravageur du soja aux USA.La consommation en poids frais (fw) a augmenté presque 2 fois quand le régime artificiel a été dilué progressivement avec de l'eau (65, 79, 86 ou 89% d'eau); le poids sec (dw) et le taux de consommation relative (RCR) ont diminué néanmoins avec la dilution. L'efficacité de digestion et d'assimilation de l'aliment consommé (digestibilité approchée, AD) a augmenté pour les trois dilutions, l'efficacité de conversion en biomasse de l'aliment digéré (ECD) a augmenté dans les dilutions à 79 et 85%, mais a diminué pour les régimes plus dilués. Par conséquent, le gain dw et le RGR, — produit de RCR x AD x ECD-, étaient identiques avec le régime à 79% fw à ceux de 65% fw, mais ont diminué pour les régimes plus dilués. Le taux relatif d'efficacité de consommation d'azote a diminué aussi avec la dilution, mais il a été compensé par un accroissement de l'efficacité de l'utilisation de l'azole tel, que le produit,-taux relatif d'accumulation de l'azote-, était le même pour les quatre régimes. La teneur en lipides des insectes a diminué, de 32% pour le régime sans dilution, à 13% pour le régime le plus dilué; elle a été le principal responsable de la diminution de RGR. L'accroissement de la consommation fw et de AD, tout en n'empêchant pas la diminution de RGR pour les deux régimes les plus dilués, a limité l'effet de la dilution (sans ces accroissements, le RGR du régime le plus dilué n'aurait été que 43% du RGR obtenu).Ces résultats indiquent que la consommation et l'utilisation d'aliments constituent un processus dynamique, et que les chenilles de A. gemmatalis comme beaucoup d'autres insectes, présentent des réactions compensatrices au changement de qualité de l'aliment. De telles variations dans la consommation d'aliments ont des conséquences écologiques (consommation accrue de substances allélochimiques potentiellement toxiques et exposition accrue aux causes biologiques de mortalité pendant l'alimentation), et intéressent l'organisation de la protection contre les insectes:-en permettant la manipulation de l'alimentation des insectes pour augmenter la consommation à court terme d'agents bioactifs, et, par ce moyen, réduire la consommation totale,-en permettant d'améliorer la modélisation des dégâts alimentaires prévisibles, qui, pour être efficace, devrait inclure les variations de l'alimentation en fonction de la qualité alimentaire de la plante cultiveé.