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.     

Food utilization by insects: Interpretation of observed differences between dry weight and energy efficiencies

Over 80% of the values of approximate digestibility (AD), efficiency of conversion of assimilated food to biomass (ECD) and efficiency of conversion of ingested food (ECI) calculated using energy terms are greater than the corresponding dry weight (DW) values, based on data for over 65 species (38 studies; number of comparative values: AD=139, ECD=128 and ECI=169). Largest positive differences (energy > DW values) are 30 (AD, ECD) and 24 (ECI) percentage points and largest negative differences (energy < DW values) are 9 (AD), 11 (ECD) and 8 (ECI) percentage points. These differences generally are least for ECI (71% of the differences fall between 0 and +5 percentage points), and AD (68%), followed by ECD (only 47% fall between 0 and +5), and they may vary with temperature, food and other factors. The differences tend to increase (esp. for ECD and ECI) when comparing later with earlier instars. Energy > DW efficiency values are commonly expected for AD because of the generally greater energy content of food than feces, and for ECD and ECI because of the generally greater energy content of insect biomass than ingested and assimilated food. Deviations from predicted differences in surveyed literature data are discussed in terms of possible methodological sources of error.

---

Résumé Plus de 80% des valeurs de la digestibilité approchée (AD), de l'efficacité de la conversion de la nourriture assimilée en biomasse (ECD) et de l'efficacité de la conversion de l'aliment ingéré (ECI), calculées en termes énergétiques, et obtenus à partir de données sur 65 espèces, sont supérieures aux valeurs des poids secs correspondants (DW): 38 études; valeurs comparatives: AD=139, ECD=128, ECI=169. Les plus importantes différences positive (énergie>valuers DW) sont de 30 (AD, ECD) et de 24 (ECI) centièmes (les différences négatives les plus fortes = 9 (AD), 11 (ECD) et 8 (ECI); ces différences sont moindres pour ECI (71% des différences tombent à 0 et +5 centièmes), et AD (68%), suivi de ECD (seulement 47% tombent entre 0 et +5). Ces différences peuvent varier avec la température, l'alimentation et d'autres facteurs; les différences tendent à croître (particulièrement pour ECD et ECI) quant on les compare plus tard avec des stades plus précoces. Energie > aux valeurs d'efficacité DW sont généralement attendues pour AD par suite du contenu énergétique supérieur de l'aliment à celui des excréments, et pour ECD et ECI par suite du contenu énergétique généralement plus élevé pour la biomasse de l'insecte que pour l'aliment ingéré et assimilé. Les écarts par rapport aux différences prédites dans les données de la littérature examinée sont analysées en considérant les sources possibles d'erreurs méthodologiques.

     

Effects of temperature on growth and efficiency of food utilization in fifth-instar caterpillars of the tobacco hornworm, Manduca sexta

Fifth-instar larvae of Manduca sexta were reared from hatching on artificial diet at 15, 20, 25, 30 and 35°C. Total development time decreased with increasing temperature. Very few larvae (12%) survived at 15°C, so this temperature was not considered further. There was some mortality at 30°C (11%), and at 35°C (50%).The absolute rate of growth in the fifth instar was faster at 25 than at 20°C, but was similar at 25, 30 and 35°C. This was true both for caterpillars that were chronically exposed to experimental temperatures (i.e. since hatching) and for those acutely exposed (i.e. reared up to fifth instar at 25°C).There was a progressive decrease with higher rearing temperatures in both the initial and final sizes of chronically exposed fifth-instar larvae. Acutely exposed caterpillars matched for initial size showed smaller temperature related differences in final size. Because of these size differences there were differences in relative growth rate which did not reflect true differences in absolute growth rate.Total food consumed by chronically exposed caterpillars was greatest at the lowest temperature (20°C), and decreased progressively with increasing temperature. The absolute rate of food consumption increased from 20 to 25°C, but did not vary significantly between 25 and 35°C. Differences in the sizes of the insects at the different temperatures meant that there were differences among relative measures of consumption that did not reflect absolute food consumption.For chronically exposed caterpillars, none of the three usual indices of food conversion efficiency (AD, ECI and ECD) varied significantly with temperature between 20 and 35°C. This implies that the effects of temperature on metabolic costs are closely matched to food consumption.Oxygen consumption increased with temperature between 20 and 25°C but was temperature compensated between 25 and 35°C.These findings are discussed in terms of their implications for the optimal temperature for growth in Manduca.     

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.     

Diet-switching by gypsy moth: effects of diet nitrogen history vs. switching on growth, consumption, and food utilization

Gypsy moth (Lymantria dispar (L.) (Lepidoptera: Lymantriidae)) larvae were reared from hatch on 1.25% N or 3.5% N artificial diet (previous diet) and switched reciprocally to the other diet (current diet) after molting into the second, third, fourth, or fifth instar. The nitrogen concentration of food consumed during previous instars had a strong residual effect on the growth rate in subsequent instars when a diet switch was made during instars two through four, but did not affect growth rate of fifth-instar larvae despite effects on food consumption and utilization. In early instars, larvae reared on 1.25% N artificial diet and then switched to 3.75% N diet had lower mass-adjusted growth rates than larvae continuously reared on 3.75% N diet. Conversely, larvae reared on 3.75% N diet and switched to 1.25% N had higher mass-adjusted growth rates than larvae reared continuously on 1.25% N diet. Relative to larvae previously reared on 1.25% N diet, fifth-instar male larvae previously reared on 3.75% N diet had slightly lower consumption rates, higher net growth efficiency (ECD), and higher gross growth efficiency (ECI). Larvae previously reared on 3.75% N diet tended to have lower food assimilation efficiency (AD) and lower nitrogen assimilation efficiency (AD(N)). Although both previous and current diet nitrogen concentration strongly affected larval growth and food utilization, the interaction term between these was not significant for any response variables except ECD and ECI. Because the interaction term reflects the effect of switching per se, the results indicate that there was a metabolic cost associated with switching, but no inherent net cost or benefit of diet-switching to growth.     

Influence of various host plants on the consumption and utilization of food by Pieris brassicae (Linn.)

Pieris brassicae (Linn.) is a destructive cosmopolitan pest of cruciferous crops. It is present wherever its host plants occur, and it is considered to be one of the most widely distributed of all the Lepidoptera. We investigated the affect of various host plants on the food consumption and utilization by P. brassicae. We quantified consumption of food, larval duration, pupal duration and weight on cabbage (Brassica oleracea var. capitata), cauliflower (Brassica oleracea var. botrytis), radish (Raphanus sativus), broccoli (Brassica oleracea var. italica) and mustard (Brassica campestris) under laboratory conditions. Insect-host relationships can be better understood by knowing the rate of food consumption, its digestibility and conversion of food eaten to body tissue. The consumption of food generally increased with the advancement of larval age. In our study we found that consumption of food was highest on radish and lowest on broccoli. The highest consumption of a particular host does not always indicate greater suitability of that host, until and unless other factors like consumption index (CI), relative growth rate (RGR), efficiency of conversion of ingested food (ECI), approximate digestibility (AD) and efficiency of conversion of digested food (ECD) are also considered. In the current investigation, factors like CI, RGR, ECI and ECD were highest on cabbage. Low body weight of pupa is associated with rapid development. On cabbage, the weight of pupa of both sexes was found lowest. Thus, from the present study, it can be concluded that cabbage is a more suitable host for P. brassicae than other host plants evaluated. Hence, on cabbage, the values of Waldbauer indices were highest and P. brassicae developed with a faster rate.     

Influence of whorl region from resistant and susceptible corn genotypes on fall armyworm (Lepidoptera: Noctuidae) growth and development

The effect of diets prepared from whorl tissue of resistant and susceptible corn genotypes, Zea mays L., on the larval growth, development, and physiology of fall armyworm, Spodoptera frugiperda (J. E. Smith), was analyzed. Larvae reared on an optimized artificial diet had a higher growth rate and developed faster than those reared on lyophilized whorl tissue from resistant and susceptible genotypes. Larvae reared on the resistant material were smaller and had a longer developmental period. Larvae reared on yellow-green and green whorl sections from resistant plants were significantly smaller than those reared on the same sections of susceptible plants. There was no significant difference in weight when larvae were reared on the yellow whorl regions from either resistant or susceptible lines. Physiological indices were determined for larvae fed resistant and susceptible lyophilized and fresh whorl material. Larvae fed resistant lyophilized material had significantly lower growth rate (GW) and efficiency of conversion of ingested food to body substance (ECI) than those reared on artificial diet or susceptible material. However, there were no significant differences in consumption index (CI), approximate digestibility (AD) and efficiency of conversion of digested food to body substance (ECD) between larvae reared on lyophilized tissue from resistant and susceptible genotypes. Larvae reared on fresh yellow-green whorl sections from resistant plants had significantly lower GW, ECI, and ECD than those reared on susceptible material. In contrast, no significant differences in any of the estimated food consumption and utilization indices were observed between larvae reared on fresh yellow whorl sections from resistant or susceptible plants. These results suggest that some components of whorls from resistant plants, especially the yellow-green region, inhibit food utilization in fall armyworm larvae.     

Legacy effects of low N diets on a vernal folivore: deprivation leads to gluttony

We tested for legacy effects of low-N diets offered to newly emerged second-instar spruce budworm (Choristoneura fumiferana) larvae for a duration of either one or two full instars on their growth and nutritional physiology in the sixth instar. The experimental design evaluated the effects of initial diet, final diet, and sex on energy consumption, assimilation, retention, and growth rates. Legacy effects were apparent after two instars of low-N diet exposure and were manifested by elevated ( approximately 10%) consumption rates (RCRs) coupled to elevated ( approximately 10%) growth rates (RGRs) and elevated ( approximately 3%) body energy densities, i.e., heightened fat deposition. However, initial dietary N levels had no legacy main effects on food assimilation efficiencies (ADs), and gross (ECI) and net (ECD) food conversion efficiencies. RCR and AD were dependent on an initial x final diet interaction (i.e., nonlinear legacy effects). RGR depended on an initial diet x sex interaction but not on an initial x final diet interaction. Therefore, the legacy effects of low-N initial diets on RGR and body energy density were simply additive to final diet effects. Final diet universally affected all indices and interacted with sex. Low-N final diets increased RCR ( approximately 41%) and decreased AD (14-18%) but unexpectedly increased ECD (21-24%) and RGR ( approximately 36%). Females generally had higher performance than males on the low-N diets but often only matched males on the high-N diets. Low-N initial diets extended larval development times ( approximately 7-26%) and lowered growth rates (6-24%) to the sixth instar, depending on duration of diet exposure, but did not affect total growth achieved by the start of the sixth instar.     

Food,feeding rates and assimilation in the land snail Cepaea nemoralis L.

Summary The natural diet of a population of Cepaea nemoralis living in a sand dune system in southwest England was investigated by field observations and examination of crop and faeces contents. The food consisted of dead material from the commoner plants on the site. Very little green matter was eaten and there was little evidence for seasonal variation in the diet.Consumption and egestion rates were measured gravimetrically on natural foods (dead leaves of Taraxacum officinale and Ononis repens) and artificial foods. Consumption rates were temperature dependent over the measured range of 5–20°C and also varied with the food substrate, an artificial food (rat food pellets) being consumed at the highest rate. High variability in the results can probably be linked to the live weight and activity cycles of the snails.Assimilation was estimated as the difference between consumption and egestion, since a trial of the ash-ratio method proved unsuitable with this animal. Assimilation efficiency did not vary with temperature over the range studied, but was dependent on food substrate, the two artificial foods (rat food pellets and lettuce leaves) being assimilated more efficiently (71–73%) than natural foods (30–44%).Faeces produced from the digestive gland could be distinguished and were measured separately. The amount produced was very small compared to the gut faeces, but increased significantly when a relatively indigestible food was used.     

Influence of parasitism byEucelatoria bryani [Dip.: Tachinidae] on the consumption and utilisation of chickpea flour diet byHeliothis armigera [Lep.: Noctuidae]

Fourth and 5th instar larvae ofHeliothis armigera (Hübner) parasitised byEucelatoria bryani Sabrosky consumed significantly less chickpea flour diet than unparasitised larvae of the same age in the laboratory. Significant reduction in diet consumption, larval weight gain and frass produced at 24 and 48 h following parasitisation was observed in 5th instar host larvae. Parasitised larvae retained a greater percentage of food ingested (AD) than did the unparasitised ones. Unparasitised 5th instar larvae ofH. armigera were more efficient in converting the ingested food (ECI) and digested food (ECD) into body substance than larvae of similar age parasitised byE. bryani.     

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.     

Using nutritional indices to study LOX3-dependent insect resistance

Induced resistance to biotic attackers is thought to be mediated by responses elicited by jasmonic acid (JA), a subset of which are lipoxygenase 3 (LOX3) dependent. To understand the importance of LOX3-mediated insect resistance, we analysed the performance of Manduca sexta larvae on wild-type (WT) and on isogenic Nicotiana attenuata plants silenced in NaLOX3 expression and JA signalling, and we used Waldbauer nutritional indices to measure the pre- and post-ingestive effects. LOX3-mediated defenses reduced larval growth, consumption and frass production. These defenses reduced how efficiently late-instar larvae converted digested food to body mass (ECD). In contrast, LOX3-mediated defenses decreased approximate digestibility (AD) in early instar larvae without affecting the ECD and total food consumption. Larvae of all instars feeding on defended WT plants behaviourally compensate for their reduced body mass by consuming more food per unit of body mass gain. We suggest that larvae feeding on plants silenced in NaLOX3 expression (as-lox) initially increase their AD, which in turn enables them to consume more food in the later stages and consequently, to increase their ECD and efficiency of conversion of ingested food (ECI). We conclude that N. attenuata's oxylipin-mediated defenses are important for resisting attack from M. sexta larvae, and that Waldbauer nutritional assays reveal behavioural and physiological counter responses of insects to these plant defenses.     

Coprophagy: a supplementary food source for two freshwater gastropods?

1. The freshwater pulmonate snail Radix peregra voluntarily and regularly fed on its faeces, whereas Bithynia tentaculata (Prosobranchia) did not.
2. With high-quality faeces as food (faeces with C : N-values below ten) Radix could grow and survived for 11 weeks. Growth was 24–30% of that achieved with control food (lettuce, Chlamydomonas and Tetramin), and body mass was close to the expected value.
3. As with real food, the percentage organic content of faeces was reduced during gut passage, with the exception of very low-quality faeces.
4. Faeces contained living cells of green algae ( Chlamydomonas reinhardii ) and diatoms ( Achnanthes lanceolata, Eunotia pectinalis ), which survived the gut passage of R. peregra and B. tentaculata . Faecal material was also rich in bacteria, with up to 150 × 10⁶ cells mg^–1 dry mass. Bacteria increased in abundance in faeces which had been evacuated for 7 days or more.
5. A further degradation of partly digested food in the faeces was indicated by the activity of the extracellular digestive enzymes cellobiase and chitobiase. These enzymes hydrolysed material with an activity of 0.2–14.5 pkat mg^–1 faecal dry mass in 1–12-day-old faeces. In faeces of Bithynia fed control food or incubated sycamore leaves, chitobiase activity was correlated with bacterial abundance.
6. Coprophagy is considered to be a suitable strategy for further degradation and utilization of refractory material for which one gut passage is too short and/or too inefficient.     

Simultaneous effects of potassium, rutin and temperature on performance ofManduca sexta caterpillars

the simultaneous effects on an insect herbivore (third instar tobacco hornwormManduca sexta (L.): Sphingidae) of temperature (daytime temperatures of 20 °C, 25 °C and 30 °C), a mineral that may play a role in plant defense (potassium) and a common allelochemical (rutin) were examined in a factorial experiment. To manipulate potassium levels, a modified diet with limited plant material was used as the base and KCl and rutin added. Temperature affected efficiency of conversion of ingested food (ECI), efficiency of conversion of digested food (ECD), time to head-capsule slippage, stadium duration, relative consumption rate (RCR) and relative growth rate (RGR) but not food consumed, biomass gained and approximate digestibility (AD). Potassium concentration influenced all of the variables except AD, time to head-capsule slippage (HCS), duration of the stadium and percent of stadium time to HCS. Rutin impacted negatively on all of the variables except food consumed. Compared to larvae on the non-rutin diets, fewer larvae fed rutin survived through molt initiation to ecdysis and fewer successfully completed ecdysis. Temperature and rutin had interactive effects for AD, ECD, RCR, RGR, time to HCS, and percent of stadium required to reach HCS. Rutin and potassium had interactive effects for biomass gained, RCR, ECI, time to HCS, duration of stadium, and percent of stadium required to reach HCS. Comparison of larval responses on an average potassium concentration (3.1%) versus high concentration (6.1%) showed that at the low daytime temperature increasing potassium concentration depressed biomass gained, but at the warmer temperatures potassium concentration had little effect unless rutin was present. In addition, potassium concentration had little impact on ECI unless rutin was present. These results indicate that significant interactive effects occur among temperature, potassium and rutin, and thus suggest that such interactive effects on larval performance may be common under field conditions, which are characterized by varying temperature and different concentrations of minerals and allelochemicals in hostplants.     

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.     

Food utilization and metabolic efficiency in larval and adult house crickets

The most rapid growth of the fat body, ovaries and residual body occurred only at the time of maximal feeding, which was the time of highest metabolic rate. Larvae ceased feeding before the next moult and lost weight. Adult females did not cease feeding and never lost weight. Mated females maintained a high feeding rate that corresponded to a constant oviposition rate with no gonotrophic cycles. Virgin females maintained a lower feeding rate and maintained fully mature eggs in a viable state for life.Growth during the last larval stadium was entirely somatic and characterized by a great accumulation of a lipid rich (65%) fat body. Net growth during the first 10 days of adult life was entirely gonodal, because the small amount of residual body growth was matched by the amount of fat body loss. However, since the residual body was 60% protein and the fat body was 60% lipid, some of the somatic protein growth had to come from the diet and some of the fat body lipid went for ovarial growth.The 20% dietary protein was more than sufficient for all somatic and ovarial growth, and some protein was catabolized for energy. The amount of uric acid in larval faeces accounted for almost all protein catabolism, but in adult females half of the catabolized protein nitrogen did not appear in the faeces. With only 5% dietary lipid both larvae and adults had to synthesize additional lipids from carbohydrates to supplement absorbed lipids for growth demands. Almost all absorbed and synthesized lipids were used for somatic growth in the larvae and for ovarial growth in the adults, because the per cent of food calories used for growth greatly exceeded the per cent dry weight of food used for growth. Based on the calories and dry weight of available fuel, almost all energy production was based on carbohydrate oxidation. Some carbohydrate was used for lipid synthesis but very little was used for growth.Probably the absorption efficiency (72%) and growth efficiency (28%) was the same for both the last instar larvae and virgin females because they ate exactly the same food. However, the larval metabolic efficiency (42%) was higher than that of females (37%), which indicated that more of the absorbed food was converted to tissue during larval growth (somatic) than during adult growth (ovarial).Of the dry weight of food eaten during the last larval stadium, 30% was egested, 32% was oxidized, 28% appeared as growth, and 10% was condensed to lipid and became part of the 28% growth. Of the dry weight of food eaten during the first 10 days of adult life, 27% was egested, 41% was oxidized, 25% appeared as growth, and 6% was condensed to lipid.     

FOOD SELECTION IN THE LAND SNAIL LIMICOLARIA FLAMMEA MULLER (PULMONATA: ACHATINIDAE)

Food consumption by juvenile and adult specimens Limicolariaflammea was measured as dry weight of food eaten. The snailwas fed weekly for 15 months on four types of food (lettuce,potato, apple and carrot) and chalk provided, hence an opportunityto select for food. Juvenile snails showed greater selectivitydepending on whether the food was needed for somatic growthor reproduction. On the other hand, when fed exclusively onone each of the same food items and given no opportunity tochoose, the snails grew better on the same food items they selectedin the choice experiment. The absorption efficiencies (AD) werehigh, the utilisation efficiency (ECI) was 1–13% and showeddistinct peaks corresponding to egg production. Values for ECDwere 1.25–15.5%. Variation in growth rates was due mainlyto utilisation efficiencies of the four foods tested. Largeamounts of chalk were eaten during the period of somatic growthand greatly reduced during the reproductive phase of L. flammea.The significance of these findings to the snails ecology isdiscussed. (Received 9 April 1991; accepted 11 June 1991)     

Effect of dietary nitrogen on gypsy moth larval nutritional indices, development and lipid content

Nutritional indices, development rates, percent dry weights and total lipids were determined in gypsy moth larvae (Lymantria dispar L.) reared on a high wheat germ (HWG) diet or diets prepared from lyophilized, ball-milled oak or pine foliage as the only source of dietary nitrogen (N). With regard to both total and proteinaceous N content, HWG diet>oak diet>pine diet. All nutritional indices measured were significantly lower in second instars fed pine diet vs. oak diet. Protein supplementation of pine diet with either casein or ovalbumin to bring total N up to the level present in oak diet resulted in small increased in approximate digestibility (AD) and effciency of conversion of ingested food (ECI), but relative growth rate (RGR) remained unaffected. The low RGR of larvae fed pine diet (unsupplemented or protein supplemented), as compared to those fed HWG or oak diet, was accompanied by significantly lower larval percent dry weight and percent total lipid. In contrast, RGR, larval percent dry weight and total lipid values were comparable in second instars fed HWG or oak diet. Insects reared from the first through the final instar on oak diet exhibited lower pupal weights compared to those reared on HWG. Casein addition to oak diet generally resulted in even more extended larval development times and further reduced pupal weights, but wheat germ addition to oak diet did not alter development rates and caused an increase in pupal weights.     

Elevated CO2 and temperature alter development and food utilization of Spodoptera litura fed on resistant soybean

Effects of elevated atmospheric CO₂ (elevated CO₂ vs. ambient CO₂) and temperature (+0.67–0.79°C vs. ambient temperature) on the developmental life cycle of Spodoptera litura and the food utilization of the fourth‐instar larvae fed on soybean (resistant cultivar Lamar vs. susceptible landrace JLNMH) grown in open‐top chambers were studied from 2013 to 2015. The results indicated that: (i) compared with ambient CO₂, elevated CO₂ significantly prolonged the duration of larva and pupa, and adult longevity; significantly decreased the pupation rate, pupal weight, fecundity, the relative growth rate (RGR), efficiency of conversion of ingested food (ECI) and efficiency of conversion of digested food (ECD); and increased the relative consumption rate (RCR) and approximate digestibility (AD). (ii) Compared with ambient temperature, elevated temperature significantly shortened the duration of larva and pupa; significantly decreased the pupal weight; and increased the RGR, RCR, ECD and ECI. (iii) Compared with the susceptible soybean accession JLNMH, the resistant soybean cultivar Lamar significantly prolonged the duration of larva and pupa; significantly decreased the pupation rate, pupal weight, adult longevity, fecundity and RGR, RCR and AD; and increased the indexes of ECD. (iv) At elevated temperature, S. litura fed on resistant vs. susceptible cultivars showed opposite trends in the RGR, RCR, AD, ECD and ECI. In addition, elevated temperature under elevated CO₂ significantly decreased the RGR (2014), ECD (2013 & 2014) and ECI (2013) and increased the AD (2013 & 2014) compared with other treatment combinations when S. litura fed on Lamar. Future climatic change of temperature and CO₂ concentration would likely affect growth and food utilization of S. litura, with increased food intake, but the reduced fecundity may compensate for the increased food consumption, resulting in no significant reduction in insect‐induced yield loss in soybean production. Nevertheless, use of insect resistant soybean cultivars will aid in ecological management of S. litura and reduce the insecticide load in soybean production.     

Optimization of gut structure and diet for higher vertebrate herbivores.

A generalized herbivore gut is modelled as (i) a well-stirred anterior chamber in which microbial fermentation occurs; (ii) a tubular reactor in which digestion but no fermentation occurs; and (iii) a posterior fermentation chamber. The rate at which the herbivore gains metabolizable energy is calculated for diets that can be eaten at different rates and contain different energy densities of easily digested cell contents, and of cell wall materials that can be fermented but not digested. The optimum gut structure for each diet is determined. Chewing probably speeds digestion and fermentation but reduces eating time. Optimal chewing times are determined for particular diets and guts. Herbivores often have a choice between poorer food that can be eaten fast and richer food that can only be eaten more slowly. Energy costs may be incurred in travelling between patches of the richer food. Optimal diet choices are predicted for herbivores with particular gut structures.