Interactions of allelochemicals with dietary constituents: effects on deterrency

Abstract. That variation in the water and nutrient content of plant tissues affects allelochemical toxicity to insects is well established. However, little is known about how these dietary constituents influence allelochemical deterrency. In vertebrates, deterrency of particular allelochemicals increases with dietary water, and decreases with an increase in dietary nutrients. We determined if these findings were relevant to phytophagous insects through an experimental design that allowed us to vary independently the content of water (70–90% fresh mass, fm, with nutrient level at 10% fm) and nutrients (10–30% fm with water level at 70% or 80% fm) in an artificial diet through use of alphacel, a non-nutritive cellulose fibre. We examined the effect of these dietary manipulations on allelochemical deterrency by comparing larval feeding responses of two noctuid species, the oligophagous Anticarsia gemmatalis Hübner and the polyphagous Spodoptera frugiperda (J. E. Smith), to a control diet and an allelochemical-treated diet in two consecutive, no-choice tests. These tests were restricted to 3 min to minimize post-ingestive influences. From an initial twelve compounds tested, all but two were excluded for the following reasons: (i) failure to elicit an intermediate level of deterrency at <1% fm (i.e. albizziin, amygdalin, hordenine, ouabain, pipecolic acid, salicin, sinigrin and umbelliferone); (ii) an apparently rapid toxic effect (nicotine hydrogen tartrate); and (iii) adsorption to alphacel (quinine hydrochloride), which may have reduced deterrency. The deterrency of caffeine and linamarin increased with dietary water but was unaffected by nutrient content for both species. The similar results for an alkaloid and a cyanogenic glycoside, with two species differing considerably in feeding habits, suggest that dietary water is likely to influence the defensive efficacy of a broader range of deterrent allelochemicals to a variety of plant-feeding insects.     

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.     

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.     

THE INFLUENCE OF DIETARY α‐SOLANINE ON THE WAXMOTH Galleria mellonella L

Plant allelochemicals are nonnutritional chemicals that interfere with the biology of herbivores. We posed the hypothesis that ingestion of a glycoalkaloid allelochemical, α‐solanine, impairs biological parameters of greater wax moths Galleria mellonella. To test this idea, we reared wax moths on artificial diets with 0.015, 0.15, or 1.5 mg/100 g diet of α‐solanine. Addition of α‐solanine to the diet affected survival of seventh‐instar larvae, pupae, and adults; and female fecundity and fertility. The diet containing the highest α‐solanine concentration led to decreased survivorship, fecundity, and fertility. The diets supplemented with α‐solanine led to increased malondialdehyde and protein carbonyl contents in midgut and fat body and the effect was dose‐dependent. Dietary α‐solanine led to increased midgut glutathione S‐transferase activity and to decreased fat body glutathione S‐transferase activitiy. We infer from these findings that α‐solanine influences life history parameters and antioxidative enzyme activities in the midgut and fat body of G. mellonella.     

Effects of chlorogenic acid- and tomatine-fed caterpillars on performance of an insect predator

Two diet experiments addressed the effects of allelochemical-fed prey (Manduca sexta caterpillars), temperature, and gender on performance of the insect predator, Podisus maculiventris. Two of the major allelochemicals in tomato were used: chlorogenic acid and tomatine. Predator performance was negatively affected by both chlorogenic acid-fed and tomatine fed-prey, and there were allelochemical by thermal regime interactions for both. Relative consumption rate and growth rate decreased at the higher levels of tomatine at the warmer thermal regime (summer conditions) but were unaffected at the cooler thermal regime (spring conditions). At the cooler thermal regime, stadium duration was prolonged when the predators were given chlorogenic acid-fed prey, but at the warmer thermal regime there was no such effect. There were several effects of gender: biomass gained, food consumed, relative growth rate and efficiency of conversion of ingested food to biomass were higher for females than males. Furthermore, the effects of thermal regime and tomatine on food consumption and biomass gained differed for females and males. In general, the hypothesis that generalist insect predators may be a selective pressure shaping host plant range of insect herbivores was supported by these results. But the occurrence of allelochemical by thermal regime interactions means that it will be difficult to determine the relative importance of plant chemistry versus predators on patterns of feeding specialization by herbivores without taking into account a third factor, temperature. Received: 20 March 1995 /Accepted: 2 August 1996     

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.     

Behavioral and morphological responses of an insect herbivore to low nutrient quality are inhibited by plant chemical defenses

Animals have several strategies to contend with nutritionally poor diets, including compensatory consumption and enhanced food utilization efficiencies. Plants produce a diversity of defense compounds that affect the ability of herbivores to utilize these strategies in response to variation in food nutritional quality. Little is known, however, about effects of allelochemicals on herbivores utilizing integrated behavioral and morphological responses to reduced food quality. Our objectives were to (1) examine how variation in diet nutritional quality influences compensatory responses of a generalist insect herbivore, and (2) determine how plant defenses affect these processes. Gypsy moth (Lymantria dispar) larvae were administered one of nine combinations of diet having low, moderate, or high nutritional quality and 0, 2, or 4 % purified aspen (Populus tremuloides) salicinoids. We quantified larval growth, consumption, frass production, and biomass allocation to midgut tissue over a 4-day bioassay. In the absence of salicinoids, larvae compensated for reduced nutritional quality and maintained similar growth across all diets through increased consumption, altered midgut biomass allocation, and improved processing efficiencies. Dietary salicinoids reduced larval consumption, midgut biomass allocation, digestive efficiencies, and growth at all nutritional levels, but the effect size was more pronounced when larvae were fed nutritionally suboptimal diets. Our findings demonstrate that integrated behavioral and morphological compensatory responses to reduced food quality are affected by plant defenses, ultimately limiting compensatory responses and reducing larval performance.     

Interactive effects of rutin and constant versus alternating temperatures on performance ofManduca sexta caterpillars

The effects of different concentrations of rutin and constant temperature (20 °C) versus alternating temperatures (23∶15 °C) on growth, molting and food utilization efficiencies of third instar tobacco hornworms (Manduca sexta) were determined. Relative consumption rate (RCR) and relative growth rate (RGR) were significantly higher for larvae at the alternating thermal regime compared to those at the constant (representing the average) temperature. With increasing concentrations of rutin, the negative effect of rutin on RCR and RGR increased for the larvae in the alternating thermal regime; however, at the constant temperature, rutin had little effect. The alternating thermal regime promoted synchrony in the timing of spiracle apolysis (the earliest morphological marker of molt). Rutin disrupted that synchrony. I discuss how patterns of host plant resistance may be altered with a decrease, in amplitude of diurnal temperatures (as has been documented recently for temperate regions) through the uncoupling of herbivore performance and allelochemical concentration. I conclude that simultaneous consideration of fluctuating temperatures and allelochemicals is advisable when assessing the effects of temperature and allelochemicals on performance of insect herbivores because interactive effects between temperature and dietary components occur and perhaps are common.     

Food consumption, utilization and detoxification enzyme activity of larvae of three polyphagous noctuid moth species when fed the botanical insecticide rotenone

We assessed the effects of the isoflavonoid rotenone, an insecticidal allelochemical occurring in various legumes, on larval performance of three polyphagous noctuid species: the corn earworm (CEW), the fall armyworm (FAW) and the southern armyworm (SAW). As rotenone concentration was increased up to 1% fresh mass in an artificial diet, neither mortality (for all three species) nor food consumption (for SAW and FAW) was significantly affected, but developmental time of the latter two species was prolonged. In contrast, for CEW developmental time was shortened and food consumption declined, especially at the two highest rotenone concentrations. Final biomass of all three species declined as dietary rotenone increased.Analysis of covariance (ANCOVA) was used to evaluate larval food utilization analogous to, but without the potential statistical problems of, the traditional ratio-based indices approximate digestibility (AD) and efficiency of conversion of digested food (ECD). Frass output, statistically adjusted to account for differences in consumption, increased with rotenone concentration for all three species, suggesting that, when ingested, this allelochemical interfered with the digestion and/or absorption of ingested food, especially at the higher concentrations tested.Significant statistical interactions in the ANCOVA necessitated the use of utilization plots to examine the linear regressions between biomass gain and absorption at each rotenone concentration. At the two highest concentrations, rotenone tended to reduce the amount of food absorbed by the larvae, as well as their ability to convert the absorbed food to biomass.When piperonyl butoxide, an inhibitor of the polysubstrate monooxygenase (PSMO) system known to metabolize rotenone, was added to the diet along with 0.5 or 1% rotenone, mortality increased significantly only for FAW, approaching or exceeding 50%. The only apparent effects of dietary rotenone on PSMO (aldrin epoxidase) activity (adjusted by ANCOVA for the covariate, crude homogenate protein) were for CEW, where activity decreased in all rotenone treatments compared to the control, and for SAW, where activity at the 3 lowest rotenone concentrations (0.001–0.1% fm) declined about 50% below that of larvae on the control diet, whereas at 0.5 and 1% rotenone, it increased approximately 2-fold over that on the control diet.     

Growth versus molting time of caterpillars as a function of temperature,nutrient concentration and the phenolic rutin

Summary A factorial experiment tested the effects of varying nutrient concentration (normal versus diluted), presence or absence of the phenolic allelochemical rutin and daytime temperature (20, 25 and 30° C) on growth, molting and food utilization efficiencies of tobacco hornworms (Manduca sexta). Two of the utilization efficiencies (approximate digestibility and efficiency of conversion of ingested food) were unaffected by temperature; the third one, efficiency of conversion of digested food, was affected by temperature but there was no consistent effect. Lower temperatures significantly increased the proportion of the stadium spent molting, with larvae at a daytime temperature of 20° C spending 9% more of the stadium in molting than larvae at 30° C. Growth time was not influenced by nutrient concentration. When temperature was low and rutin absent, molt time and the proportion of the stadium spent molting were affected by nutrient concentration. Addition of the phenolic rutin did not have an appreciable effect on growth time or digestive processes. However, it increased molting time by 7 to 14% and thus increased the duration of the stadium and reduced relative growth rate. These results indicate that the effect of food quality on growth rate is a function of the thermal conditions of insect herbivores. At cooler temperatures, a disproportionate increase in time spent molting, rather than altered food utilization efficiencies, contributed to lower growth rates. The consequences for larval growth of fluctuating temperatures due to diurnal cycles and the presence of predators forcing larvae into thermally suboptimal microhabitats are discussed.     

Combined effects of night-time temperature and allelochemicals on performance of a generalist insect herbivore

To assess the pattern of temperature influencing the effect of allelochemicals on growth of insect herbivores and to examine the potential effect of warmer nights due to global warming, we examined the simultaneous effects of allelochemicals and warmer night-time temperatures on an insect herbivore (Spodoptera exigua; Lepidoptera: Noctuidae). Dietary chlorogenic acid, rutin and tomatine levels reflected those occurring naturally in the leaves of tomato, a hostplant of this herbivore. We compared the effects of four thermal regimes having a daytime temperature of 26 °C , with the night-time temperature increased from 14 to 26 °C by increments of 4 °C . The effect of a particular allelochemical on developmental rate was similar among the four thermal regimes. Chlorogenic acid and tomatine each reduced final larval weight, but there was no effect of night-time temperature. In contrast, rutin had no effect on final weight, whereas final weight declined with increasing night-time temperature. Night-time temperature did not influence amount eaten. Larvae ate less when chlorogenic acid or tomatine was in the diet. For each allelochemical, there were no allelochemical by thermal regime interactions. In addition, we compared the effects of allelochemicals and the thermal regime of 26:14 °C and constant 20 °C , which was the average temperature of the 26:14 °C regime. Developmental rate was lower at the constant 20 °C regime, chlorogenic acid and tomatine each depressed developmental rate, and there were no allelochemical by thermal regime interactions. Thus, regardless of the specific allelochemical or amount, the pattern of response at the fluctuating regime was similar to that at the constant temperature. In contrast, comparison of the thermal regime of 26:22 °C and constant 24 °C , which was the average temperature of the 26:22 °C regime, showed several allelochemical by thermal regime interactions. At the 26:22 °C regime, developmental rate was disproportionatly higher at the maximal rutin concentration compared to that at constant 24 °C . At the constant 24 °C , final larval mass was disproportionately lower at the moderate tomatine concentration compared to that at the 26:22 °C regime. Because these results differ from that of other studies examining another species, it appears that the response to incremental changes in night-time temperature will reflect the allelochemicals and insect species tested. The contrast between the constant 24 °C and 26:22 °C regimes indicates that even small fluctuations (±2 °C ) in temperature over 24 h can yield differences in the response to an allelochemical.     

Food intake, conversion efficiency, and feeding behaviour of tobacco hornworm caterpillars given artificial diet of varying nutrient and water content

ABSTRACT Fifth stadium tobacco hornworm caterpillars, Manduca sexta (L.), given artificial diet diluted to varying extents with either cellulose or water compensated for the food's reduced nutrient content by eating more of it. This compensation was, however, in most cases not sufficient to maintain normal growth rates. When the water content of the diet was reduced, the insects ate less than the usual fresh weight of food but maintained their intake of nutrients. Nevertheless, growth rate was impaired. The insects were better able to compensate for dilution of their food with water than with cellulose. The efficiency of conversion of ingested food (ECI) was decreased when the diet was adulterated with cellulose. At moderate dilution (50% nutrient) this was due mostly to decreased approximate digestibility (AD), but at greater dilution (25% and 10% nutrient content) the efficiency of conversion of digested food (ECD) was decreased. ECI was maintained when the water content of the diet was increased to give 50% nutrient concentration, but was decreased when water content was changed more radically (200%, 25% and 10% nutrient diets). This was due mostly to increased metabolic costs (decreased ECD) in all cases. The retention time of food in the gut was progressively decreased (i.e. speed of passage was increased) as nutrients were replaced by cellulose. By contrast, dilution of the diet with water resulted in only slight changes in retention time, except at extreme dilution (10% nutrient content) when retention time was reduced. Compensation of food intake was achieved by spending more (or less) time eating. Video analysis of feeding behaviour showed that there were significant changes in the length of feeding bouts and of interfeed gaps when caterpillars fed on diets of altered composition. For diets diluted with cellulose, changes in bout length and bout frequency contributed substantially to the increased time spent feeding on the adulterated food. For diets diluted with water, however, almost all of the compensatory change in behaviour was due to increased bout length, with bout frequency affected only slightly. This suggests that volumetric feedback contributes principally to the termination of feeding bouts in caterpillars, while nutrient flow may affect both the initiation and termination of feeding.     

Phytochemical defences and performance of specialist and generalist herbivores: a meta-analysis

1. Phytochemical coevolution theory, a long-standing paradigm in plant–insect interactions, predicts that specialist herbivores are less negatively affected by the allelochemicals of their host plants than are generalist herbivores. Although this theory is prevalent in plant–insect science, it is not always supported by empirical studies measuring the performance of specialist and generalist insects in response to allelochemicals. 2. The present study aimed to investigate: (i) whether there a difference between specialist and generalist performance in response to allelochemicals and (ii) whether the effect of allelochemicals on specialists and generalists depend upon allelochemical class or insect order. 3. A meta-analysis was conducted incorporating 76 effect sizes drawn from studies that directly compared the performance of specialist and generalist insects in response to treatment and control diets. Most of the effect sizes were related to the performance metric growth, the insect order Lepidoptera, and the allelochemical class nitrogen-containing compounds. 4. As predicted by phytochemical coevolution theory, specialist insects responded less negatively to allelochemicals of their hosts than generalist insects in terms of growth. There were no significant differences in terms of fecundity or survival, or among allelochemical classes or insect orders. 5. These results support the prediction of phytochemical coevolution theory that specialist insects respond less negatively to allelochemicals of their hosts than generalists, although only in terms of growth.     

Does dietary P affect feeding and performance in the mixed-feeding grasshopper (Acrididae) Melanoplus bivitattus?

Although consequences of limited dietary protein and carbohydrate to performance are well studied for terrestrial insect herbivores, the importance of phosphorus (P) remains poorly understood. We examined the significance of dietary P to performance in fifth-instar nymphs of the grasshopper Melanoplus bivittatus fed artificial diets. Consumption, digestion, developmental rate, and growth in response to different levels of P nested within standard-Protein and carbohydrate diets were determined. Developmental rate was slowest on high-P diets; protein:carbohydrate concentration and P in diets affected frass production and consumption. Approximate digestibility and conversion of digested food were primarily influenced by the protein:carbohydrate quality of the diet but not P. Mass gain was marginally lower in the low-Protein:high carbohydrate diet used in this study. At the individual level, other than small effects to developmental rate at high concentrations for M. bivittatus, dietary P otherwise seems to have little effect on nymphal performance. To the degree that it is important, effects of dietary P depend on the concentrations of protein and carbohydrate in the diet.     

Tannins — a dietary problem for hand-reared grey partridge Perdix perdix after release?

A 4-week feeding trial on 22 grey partridges Perdix perdix was conducted in this study. Seven birds were fed commercial poultry food, seven natural food and eight commercial poultry food containing 6% of quebracho-tannin. Our results suggest that 6% dietary tannin, when added to a commercial food with high protein content, effects the grey partridge only slightly. No difference was seen in food consumption and body mass remained stable. However, birds fed tannin had longer small intestines, which most probably indicate gastrointestinal detoxication. They also excreted a high amount of tannin in their faeces. In addition, no between-group variation was seen in cytochrome P450 enzymes. Birds fed natural food had high concentration of nitrogen in intestinal excreta and high plasma alanine concentrations. They also suffered a rapid decrease in body mass after the change in diet and their body mass remained low. This may indicate increased protein excretion and/or catabolism of endogenous nutrient reserves. Potential short-term effects of the change in diet were seen in plasma. These findings coincide with the high mortality period of birds released into the wild.     

Influences of maternal caffeine on the neonatal rat brains vary with the nutritional states

The potential effect of maternal caffeine ingestion upon total brain protein and the concentration of two prototype neuropeptides, thyrotropin-releasing hormone (TRH) and its derivative, cyclo (His-Pro) in neonates was examined during the nursing period in the context of variable maternal protein intake. Maternal caffeine intake (2 mg/100 g body weight) significantly increased the total brain protein of neonates derived from dams fed a 6% casein diet, but not from dams fed a 12%- or 20%-casein diet. Maternal caffeine consumption significantly increased the amount of cyclo (His-Pro) in the neonatal brains in all groups. The percent increments in pups from dams fed 6%, 12%, and 20% casein diets were respectively 137%, 131%, and 120%. By contrast, no significant alterations were observed in TRH concentrations between caffeine and control groups. It is concluded that maternal caffeine can influence neonatal brain protein and cyclo (His-Pro) during nursing under conditions of protein-energy malnutrition.     

Whole-body protein turnover reveals the cost of detoxification of secondary metabolites in a vertebrate browser

The detoxification limitation hypothesis predicts that the metabolism and biotransformation of plant secondary metabolites (PSMs) elicit a cost to herbivores. There have been many attempts to estimate these costs to mammalian herbivores in terms of energy, but this ignores what may be a more important cost—increases in protein turnover and concomitant losses of amino acids. We measured the effect of varying dietary protein concentrations on the ingestion of two PSMs (1,8 cineole—a monoterpene, and benzoic acid—an aromatic carboxylic acid) by common brushtail possums (Trichosurus vulpecula). The dietary protein concentration had a small effect on how much cineole possums ingested. In contrast, protein had a large effect on how much benzoate they ingested, especially at high dietary concentrations of benzoate. This prompted us to measure the effects of dietary protein and benzoate on whole-body protein turnover using the end-product method following an oral dose of [¹⁵N] glycine. Increasing the concentration of dietary protein in diets without PSMs improved N balance but did not influence whole-body protein turnover. In contrast, feeding benzoate in a low-protein diet pushed animals into negative N balance. The concomitant increases in the rates of whole-body protein turnover in possums eating diets with more benzoate were indicative of a protein cost of detoxification. This was about 30 % of the dietary N intake and highlights the significant effects that PSMs can have on nutrient metabolism and retention.     

Filtration and digestion responses of an elementally homeostatic consumer to changes in food quality: a predictive model

In the study of the stoichiometric relationship between autotrophs and herbivores, attention has been largely focused on effects of the encountered mismatch between needs and supplies of an element on herbivore growth and ecosystem processes. Herbivore adaptation to poor food quality has rarely been investigated. This study presents a predictive model of feeding, assimilation, digestion and excretion of Daphnia facing a dietary deficiency in phosphorus. Biochemical compounds in the food were divided into phosphorous and non-phosphorus compounds. It was assumed that Daphnia is able to differently assimilate both types of compounds by regulation of target specific digestive enzymes. Feeding rate was regulated by optimal gut residence time of food particles, and assimilation efficiency by gut residence time and optimal secretion of both classes of gut enzymes. The model predicted the optimal strategy for a consumer facing an elementally imbalanced diet: (1) increase the ingestion rate, and (2) increase the secretion rate of both classes of gut enzymes. It resulted in decreased C and nutrient assimilation efficiencies, increased C feeding costs, and reduced growth rate. Sensitivity analysis showed that these predictions were qualitatively not influenced by parameter values. An alternative model was tested that includes an additive term allowing the direct excretion of C assimilated in excess. Results showed that this strategy is not optimal for the consumer growth rate. In conclusion, the model supports the hypothesis that carbon ingested in excess may generate energy that can be used to obtain more nutrients by increased feeding rate.     

Effects of light availability on host plant chemistry and the consequences for behavior and growth of an insect herbivore

We examined how light availability influenced the defensive chemistry of tomato (Lycopersicon esculentum: Solanaceae). Tomato plants were grown either in full sunlight or under shade cloth rated at 73%. Leaves from plants grown in full sunlight were tougher, had higher concentrations of allelochemicals (chlorogenic acid, rutin and tomatine), and had less protein than leaves from plants grown in shade. We determined how these differences in host plant quality due to light availability affected the behavior and growth of a Solanaceae specialist, Manduca sexta. Both in the greenhouse and in the field, caterpillars on shade-grown plants grew heavier in a shorter amount of time than those on plants that had previously been grown in full sunlight. In contrast, the effects of previous light availability to plants on caterpillar behavior appeared to be minor.To further investigate how light availability to plants influenced herbivore growth, we examined the effects of leaf-powder diets made from tomato leaves of different ages (new, intermediate, or mature) grown in full sunlight or shade on caterpillar performance. Caterpillars fed diets made from plants grown in shade consumed less but grew faster than larvae fed diets made from tomato plants grown in full sunlight. Caterpillars fed diets made from new leaves grew larger in less time than caterpillars fed diets made from intermediate aged leaves. Caterpillars did not survive on the mature leaf powder diets. There were plant-light treatment by larval thermal regime interactions. For example, at 26:15 °C , plant-light treatment had no effect on stadium duration, but at 21:10 °C, stadium duration was prolonged with the full sunlight-new leaf diet compared with the shaded-new leaf diet. In a second diet experiment, we examined the interactive effects of protein and some tomato allelochemicals (rutin, chlorogenic acid and tomatine) on the performance of caterpillars. There were food quality by thermal regime interactions. For instance, at 26:15 °C , neither protein nor allelochemical concentration influenced stadium duration, whereas at 21:10 °C, stadium duration was prolonged with the low protein-high allelochemical diet, which simulated full sunlight leaves. In sum, light availability to plants affected defensive chemistry and protein concentration. The difference in food quality was great enough to influence the growth of a specialist insect herbivore, but the effects were temperature-dependent.     

Sagebrush and grasshopper responses to atmospheric carbon dioxide concentration

Summary Seed- and clonally-propagated plants of Big Sagebrush (Artemisia tridentata var.tridentata) were grown under atmospheric carbon dioxide regimes of 270, 350 and 650 μl l⁻¹ and fed toMelanoplus differentialis andM. sanguinipes grasshoppers. Total shrub biomass significantly increased as carbon dioxide levels increased, as did the weight and area of individual leaves. Plants grown from seed collected in a single population exhibited a 3–5 fold variation in the concentration of leaf volatile mono- and sesquiterpenes, guaianolide sesquiterpene lactones, coumarins and flavones within each CO₂ treatment. The concentration of leaf allelochemicals did not differ significantly among CO₂ treatments for these seed-propagated plants. Further, when genotypic variation was controlled by vegetative propagation, allelochemical concentrations also did not differ among carbon dioxide treatments. On the other hand, overall leaf nitrogen concentration declined significantly with elevated CO₂. Carbon accumulation was seen to dilute leaf nitrogen as the balance of leaf carbon versus nitrogen progressively increased as CO₂ growth concentration increased. Grasshopper feeding was highest on sagebrush leaves grown under 270 and 650 μl l⁻¹ CO₂, but varied widely within treatments. Leaf nitrogen concentration was an important positive factor in grasshopper relative growth but had no overall effect on consumption. Potential compensatory consumption by these generalist grasshoppers was apparently limited by the sagebrush allelochemicals. Insects with a greater ability to feed on chemically defended host plants under carbon dioxide enrichment may ultimately consume leaves with a lower nitrogen concentration but the same concentration of allelochemicals. Compensatory feeding may potentially increase the amount of dietary allelochemicals ingested for each unit of nitrogen consumed.