Food distance and its effect on nutrient balancing in a mobile insect herbivore

We investigated how the distance between foods of differing nutrient content affects macronutrient (protein and digestible carbohydrate) regulation and the patterns of food acquisition, movement and feeding activity in fifth-instar nymphs of Locusta migratoria (L.) (Orthoptera: Acrididae). We placed individual insects into one of three differently sized circular arenas (20, 40 or 80 cm diameter) that contained four dishes of chemically defined synthetic food. One of these dishes contained high-protein, low-carbohydrate food (P), and the other three dishes contained low-protein, high-carbohydrate food (C). Alone, these foods are nutritionally unbalanced, but together they are complementary. Regardless of arena size, locusts regulated their protein-carbohydrate intake to similar points, and in all three arenas they ate preferentially from the dish containing the P-food. We also recorded the patterns of foraging behaviour for 12 h on days 1 and 4 of the experiment, which allowed us to determine the effect of distance between foods and whether behaviour was modified with experience in the different arenas. Locusts' foraging behaviour in small arenas was similar on days 1 and 4, and the time they spent in different parts of the arena did not differ from that predicted by the simplest random probability model. In contrast, the foraging behaviour of locusts in the medium and large arenas changed between days 1 and 4, including spending more time in the area of the arena containing the P-food and visiting fewer of the C-food dishes. We discuss how distance between foods influences foraging strategies, and the possible role of learning in its development. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Nutritional regulation in nymphs of the German cockroach,Blattella germanica

Synthetic foods varying in protein-carbohydrate ratio and total nutrient concentration were used in food selection experiments to investigate the ingestive and post-ingestive regulation of macronutrients by male and female nymphs of the German cockroach, Blattella germanica (L.) (Dictyoptera: Blattellidae). For both nutrient imbalance and food dilution, food ingestion varied between treatments with the effect that nutrient ingestion was regulated. However, this mechanism was insufficient to compensate for some food dilution treatments. In those cases, the regulation of protein intake was prioritised over that of carbohydrate intake, and two additional regulatory responses were seen. Firstly, cellulose digestion supplemented shortfalls in dietary soluble carbohydrates, and secondly the feeding period within the stadium was prolonged. These ingestive, post-ingestive and developmental responses were orchestrated in such a way that, in all treatments, nutrient gain approached similar levels, despite the variation in food properties.     

Food mixing strategies in the desert locust: effects of phase, distance between foods, and food nutrient content

Food mixing strategies were compared in the cryptically coloured, relatively sedentary `solitarious' and the highly mobile, conspicuously coloured `gregarious' phases of the desert locust, Schistocerca gregaria. Based on phase related differences in behaviour and nutritional regulatory responses, we predicted that solitarious nymphs, compared to gregarious nymphs, would move less between nutritionally complementary foods, particularly as the distance between the foods increased. We manipulated the nutritional composition [protein (p) and digestible carbohydrate (c) content] of two foods in an experimental arena and varied the distance between the foods using a factorial experimental design. Results indicated that in general, solitarious nymphs showed greater fidelity to individual food dishes than did gregarious insects (i.e., they concentrated their feeding mainly on one dish). However, results also demonstrated that for both phases fidelity to a particular food dish increased as the distance between the dishes increased, and that the number of switches between dishes decreased with increasing distance. In the smallest arenas, though, gregarious nymphs switched more frequently between the two food dishes than solitarious nymphs, even when the two dishes contained the same, near-optimal food (p18:c24). When challenged by having the two dishes either placed furthest apart (2 m) or more divergent in nutritional composition (p29:c13 vs. p7:c35), insects of both phases regulated protein intake more strongly than carbohydrate intake, by eating more from the dish containing higher-protein food.     

Socially Influenced Behaviour and Learning in Locusts

As a part of our research on the evolution of social learning in insects, we examined socially influenced behaviour and social learning in desert locust (Schistocerca gregaria) nymphs and adults. In the nymphs, the only positive effect we documented was an increased tendency to feed while in the company of another locust than alone. The adults, on the other hand, showed significant preference for joining others (local enhancement) in both the contexts of feeding and egg laying. Neither nymphs nor adults, however, showed social learning. Our preliminary analyses pointed to locusts as a likely insect that might possess social learning. Our research, when taken together with research on phase‐shifts and swarm/marching behaviour of gregarious locusts, suggests that the behavioural dynamics of gregarious locusts may make local enhancement but not social learning beneficial. The possible difference we documented between the nymphs and adults could enable us to further explore the proximate and ultimate mechanisms that underlie socially influenced behaviour.     

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.     

Inhibiting parabrachial fatty acid amide hydrolase activity selectively increases the intake of palatable food via cannabinoid CB1 receptors

These studies investigated feeding responses to indirect activation of parabrachial cannabinoid CB1 receptors. Arachidonoyl serotonin (AA5HT), an inhibitor of the endocannabinoid degradative enzyme, fatty acid amide hydrolase (FAAH), was infused into the parabrachial nucleus of male Sprague-Dawley rats, and intakes of high-fat/sucrose pellets and standard rodent chow were subsequently evaluated under various feeding schedules. FAAH blockade stimulated the intake of high-fat/sucrose pellets that were presented daily for 4 h during the light period, with compensatory decreases in the consumption of standard chow during the ensuing 20 h. These diet-selective changes were repeated on the next day, indicating a shift in feeding toward the more palatable diet that lasted for 48 h after a single infusion. The cannabinoid CB1 receptor antagonist, AM251, blocked the orexigenic actions of AA5HT, implicating CB1 receptors in mediating the feeding responses to FAAH inactivation. When the feeding schedule was reversed, AA5HT produced nominal increases in the consumption of standard chow for the 4-h access period, but substantial increases in the intake of high-fat/sucrose during the following 20-h interval. When presented with only high-fat/sucrose diet for 24 h, AA5HT increased 24-h food intake. In contrast, when given 24-h access only to standard chow, AA5HT failed to affect intake. Therefore, indirectly activating parabrachial CB1 receptors by blocking the degradation of native ligands selectively stimulates the intake of palatable food, with differential actions on total energy intake depending upon the feeding schedule. Our results support a role for parabrachial cannabinoid mechanisms in providing physiological regulation to neural substrates modulating feeding, energy balance, and behavioral responses for natural reward.     

Behavioural correlates of phenotypic plasticity in mouthpart chemoreceptor numbers in locusts

Rearing locusts in an impoverished chemosensory environment leads to fewer chemoreceptors developing on the mouthparts and antennae as adults but the behavioural relevance of these changes remains unknown. To address this question, locusts were reared for the final two larval stadia on either a single, nutritionally near-optimal synthetic food ('plain' pretreatment), or a diet comprising two nutritionally complementary foods containing two added flavours ('mix' pretreatment). Insects reared on the 'mix' diet had a mean 20% more chemosensilla on the maxillary palps than those fed on the 'plain' diet. Following an equilibration period, when all newly moulted adults could feed on two nutritionally complementary foods, insects were food deprived for 2 or 6 h, and then given a test meal of a single balanced food at one of two dilutions whilst their behaviour was recorded. 'Mix'-pretreated locusts had a shorter latency to feed and were more likely to reject the test food upon first contact if deprived for only 2 h; but if they did take a meal it lasted longer and contained fewer pauses. Using sensilla number as a covariate removed the statistical significance of pretreatment regime, indicating that sensilla number, or some close correlate of it, can largely account for the variation in behaviour. This suggests that sensilla numbers are behaviourally relevant; particularly where locusts are not greatly food deprived and faced with marginally acceptable foods.     

Regulation of water ingestion by the locust, Chortoicetes terminifera: the effect of injections into the haemolymph

Adults of the locust Chortoicetes terminifera which had been deprived of water for 24 h took larger 'meals' of water than insects which had been deprived for 2 h. Water or dilute solutions of NaCl injected into the kaemcoel reduced the amount of water ingested by 24-h, deprived locusts. Injections of more concentrated solutions of NaCl (0.5 m) or glycerol (1.0m) increased the water intake of 2-h deprived locusts. Water injections into 24-h deprived locusts did not reduce their responsiveness to sucrose and it is concluded that haemolymph osmotic pressure plays an important role in determining the specific responsiveness of C. terminifera to water. The time course of the effects of injection indicates that the effects of haemolymph osmotic pressure on water responsiveness are mediated neurally. Injections of water into 24-h deprived insects caused a marked reduction in the sizes of their meals of seedling wheat. It is suggested that the mechanisms responsible for the regulation of water intake might limit total intake when locusts have access only to fresh vegetation of high water content.     

The relationship between plant acceptability and suitability for survival and development of the polyphagous grasshopper,Schistocerca americana (Orthoptera: Acrididae)

The feeding behavior of final-instar nymphs ofSchistocerca americana was observed when they first encountered plants that ranged in acceptability from being eaten in large amounts to not being eaten at all. Growth and survival on the same plants through the last stadium were also studied and the results combined as a suitability index to facilitate comparison with the behavior. Although the plants that were eaten most gave the highest suitability index and those that were not eaten permitted no survival, there was no simple relationship between the amounts eaten and the suitability for growth and survival. The possibility that the insects might become habituated to plants that were initially unacceptable was investigated, but no habituation was found over a 3-day period. It is suggested that food intake is largely determined by the presence of deterrent compounds in the less acceptable foods and that nutritional differences between the plants are likely to have been of minor importance. The behavior on some foods suggests that food aversion learning may be involved. It is concluded that the variability of the insects' behavior makes it impossible to predict the suitability of a plant from their immediate behavioral responses. In the field, insects may sometimes reject foods that would be suitable for survival and development, and feed on plants that are nutritionally deficient or even toxic.     

The role of food distribution and nutritional quality in behavioural phase change in the desert locust

The behaviour of herbivorous insects is influenced by their nutritional state. Nutrition-induced behavioural changes are often interpreted as adaptive mechanisms for controlling nutrient intake; however, their influence on other life history traits has received far less attention. We investigated the effect of food quality and distribution on the behaviour and phase state of desert locusts, Schistocerca gregaria Forsk?l (Orthoptera, Acrididae), which change from the 'solitarious' to the 'gregarious' phase in response to population density. Phase change involves many morphological, physiological and behavioural changes. Solitarious insects are cryptic whereas gregarious locusts aggregate. Individual phase change is stimulated by mechanical contact with other locusts. A clumped resource distribution promotes change to the gregarious phase by increasing crowding and contact between individuals. In this study, we found that the effect of food distribution on locust phase depended on the nutritional quality of the food. We used three synthetic food treatments: near optimal, dilute and a choice of two unbalanced but complementary foods. Clumped resource distribution led to increased gregarization in the dilute and the complementary diet treatments. This effect was particularly pronounced on the complementary foods, owing to the interaction of crowding and locomotion. Gregarization was most pronounced in the dilute diet treatment, owing to increased activity. These diet-induced effects are explained in terms of behavioural changes in locomotion, quiescence and feeding that are consistent with what is known from earlier work on locust feeding behaviour and behavioural phase change. Copyright 2000 The Association for the Study of Animal Behaviour.     

The selection of nutritionally balanced foods by Locusts migratoria: the interaction between food nutrients and added flavours

Abstract. To investigate the ability of insects to discriminate between nutritionally balanced and unbalanced foods, fifth-instar nymphs of Locusta migratoria L. (Orthoptera: Acrididae) were given one of thirteen pairings of artificial foods differing only in their concentration of protein and carbohydrate. One food, containing a previously estimated optimum balance of these nutrients, was common to each pairing. The relative intake of this 'target' food increased on the more extreme pairings, i.e. the insects demonstrated increasingly strong preference for the balanced food when it was paired with unbalanced alternative foods. When distinctive non-nutritional flavours (tannic acid and amygdalin) were added to one or both of the foods, the insects' intake of the unbalanced foods increased, particularly on high-protein foods. It is suggested that learned associations were not involved in the observed patterns of selection and that the added flavours disrupted the chemosensory responses to nutrients used by the insects to differentiate foods in this context.     

Compensatory dietary selection occurs in larval Locusta migratoria but not Spodoptera littoralis after a single deficient meal during ad libitum feeding

ABSTRACT Final instar nymphs of Locusta migratoria (L.) and larvae of Spodoptera littoralis (Boisduval) were given an artificial diet deficient in either protein or digestible carbohydrate for a single meal during ad libitum feeding, after which they were provided with a choice of two diets, one containing protein but no digestible carbohydrate and the other containing carbohydrate but no protein. Detailed analyses of feeding behaviour showed that locusts exhibited a degree of compensatory dietary selection following the single deficient meal. No such response was evident for the caterpillars, although previous work (Simpson et al. , 1988) has demonstrated that compensatory selection behaviour is marked in this species after periods of 4 h or more on the same deficient diets. These results show that locusts are able to respond extremely rapidly to the nutritional quality of their food by utilizing nutritional feedbacks. This capability may have important implications for the study of foraging strategies in other herbivorous insects.     

Salt Content Impacts Food Preferences and Intake among Children

Decreasing dietary sodium intake, which can be achieved by reducing salt content in food, is recommended. Salt contributes to the taste of foods and makes them more enjoyable. Whether a food is liked or disliked is an important determinant of food intake, especially among children. However, the role of salt in children''s food acceptance has received little attention. The impact of salt content on children''s hedonic rating and intake of two foods was investigated in children. Using a within-subject crossover design, we recruited 75 children (8–11 years) to participate in five lunches in their school cafeteria. The target foods were green beans and pasta. The added salt content was 0, 0.6 or 1.2 g/100 g. The children''s intake (g) of all lunch items was measured. The children provided their hedonic rating of the food, a preference ranking and a saltiness ranking in the laboratory. Children could rank the foods according to salt content, and they preferred the two saltier options. A food-specific effect of salt content on intake was observed. Compared to the intermediate level (0.6 g salt/100 g), not adding salt decreased green bean intake (−21%; p = 0.002), and increasing the salt content increased pasta intake (+24%; p<0.0001). Structural Equation Modeling was used to model the relative weights of the determinants of intake. It showed that the primary driver of food intake was the child''s hunger; the second most important factor was the child''s hedonic rating of the food, regardless of its salt content, and the last factor was the child''s preference for the particular salt content of the food. In conclusion, salt content has a positive and food-specific effect on intake; it impacted food preferences and intake differently in children. Taking into account children''s preferences for salt instead of their intake may lead to excessive added salt.     

The pattern of feeding of first instar nymphs of Schistocerca americana

ABSTRACT The feeding behaviour of nymphs Schistocerca americana (Drury) was recorded throughout the light phases of the first stadium using a behavioural event recording program on a microcomputer. Most food was ingested on day 4 of the stadium, very little on day 1 and none on day 6, the final day of the stadium. Only 26–45% of the total food intake over the stadium occurred during the dark periods and more food was eaten in the last 4 h of each light period than earlier. Feeding occupied only about 5% of the total time. Variation in food intake was a consequence primarily of changes in the numbers of meals. After day 1, the average meal size did not change significantly. Meal length increased relatively slightly compared with meal size as a consequence of an increase in the proportion of time spent feeding in each meal as meal size increased. Meal size was correlated with the length of the previous interfeed suggesting a volumetric regulation. The distribution of pauses within meals was also consistent with the thesis that meal size is governed by the level of excitation in the central nervous system at the start of the meal.     

Changes in haemolymph octopamine levels associated with food deprivation in the locust, Schistocerca gregaria

ABSTRACT. Levels of octopamine in the haemolymph and locomotor activity have been measured over 24 h in fed and food-deprived adult male locusts, Schistocerca gregaria (Forskål). Locusts are predominantly diurnal, the locomotor activity of individuals with continuous access to food was higher during the photophase. The basal level of octopamine in haemolymph sampled during the first quarter of the light cycle was 7.8±1.1 pg/μl (51 nm). Depriving locusts of food at the start of the photophase for 9 h caused an increase in both the speed of movement and total activity compared to fed individuals. There was a corresponding increase in the haemolymph octopamine concentration which continued for as long as the insects were unable to feed, reaching a concentration of 20.7 ± 3.7 pg/μl (135 nM). While declining, the levels of octopamine remained higher in fed locusts for up to 4 h after replacing the food. The results are discussed in relation to the regulation of feeding behaviour and a proposed neurohormonal role of octopamine in insects.     

A selective orexin-1 receptor antagonist reduces food consumption in male and female rats

A variety of evidence implicates the orexins, especially orexin-A, in the regulation of food intake, but it has not been established whether this effect is mediated by the orexin-1 or orexin-2 receptor. In the present study, a selective orexin-1 receptor antagonist, 1-(2-methylbenzoxazol-6-yl)-3-[1,5]naphthyridin-4-yl urea hydrochloride (SB-334867-A), was administered intraperitoneally to rats under various conditions, and food consumption was subsequently measured over 24 h. In male rats, a single dose of SB-334867-A (30 mg/kg, i.p.) given during the light phase reduced both orexin-A-induced food intake (7 nmol, i.c.v.) and feeding stimulated by an overnight fast for 4 h. When given at the start of the dark phase, food consumption was reduced in both male and female rats over 24 h. Daily injections at the start of the dark phase for 3 days reduced natural feeding in male rats over 24 h on days one and three. These findings demonstrate direct inhibition of orexin-A induced food intake with a selective orexin-1 receptor antagonist. Furthermore, the suppression of nocturnal feeding and food intake stimulated by an overnight fast supports other evidence that orexin-A is involved in the regulation of natural feeding and suggests that orexin-1 receptor antagonists could be useful in the treatment of obesity.     

The functional significance of switching interval in food mixing by Locusta migratoria

To investigate the consequences of different patterns of switching between nutritionally complementary foods, nymphs of Locusta migratoria L. (Orthoptera: Acrididae) were forced to alternate between pairs of artificial foods differing only in their protein and carbohydrate content at one of several fixed intervals. Total food and nutrient intake over 6days were very similar in all groups despite the considerable variation in temporal patterns of nutrient intake. The effects of these treatments were determined using measurements of growth, stadium duration and carcass composition. Those given a low variance food pairing showed little effect in any of the measures while those given a high variance pairing showed no differences in dry weight growth or stadium duration but did differ in protein and lipid content. There was little evidence from our measurements that an ad-libitum alternation time of 4h previously observed in a laboratory experiment was physiologically optimal, suggesting that ecological factors may underlie such behaviour. Possible compensatory mechanisms used by L. migratoria are suggested.     

Compensation by locusts for changes in dietary nutrients: behavioural mechanisms

ABSTRACT. The detailed behavioural mechanisms underlying an instance of compensation for changes in dietary nutrients are described for the first time in an insect. Nymphs of Locusta migratoria L. were given one of four artificial diets on the third day of the fifth instar, and their feeding patterns recorded in detail for 12 h. The diets represented combinations of two protein and two digestible carbohydrate levels (28% and 14% dry weight) presented in an otherwise complete nutrient mix. At the nutrient levels used, locusts regulated their intake of food with respect to protein but not digestible carbohydrate in the diet. They ate more of the lower protein diets by eating the same sized meals more frequently than insects fed on the higher protein diets. Compensation was not complete over the 12 h observation period: insects on the lower-protein diets ingested and absorbed 72% as much nitrogen as those insects fed on the higher-protein diets. Possible physiological mechanisms underlying the behavioural mechanisms are discussed.     

绿僵菌侵染后东亚飞蝗营养生理参数变化研究

本文采用饵剂饲喂方法研究东亚飞蝗Locusta migratoria manilensis(Meyen)营养生理参数变化,结果表明:绿僵菌(Metarhizium anisopliae)侵染后,5个温度梯度下蝻期总取食量分别为96.4、108.3、131.9、103.2、249.5mg;21、24、27、30℃4个温度...     

The homeostatic feeding response to fasting is under chronostatic control

ABSTRACT

Eating behavior is controlled by the energy needs of the organism. The need to provide a constant supply of energy to tissues is a homeostatic drive that adjusts feeding behavior to the energetic condition of the organism. On the other hand, food intake also shows a circadian variation synchronized to the light-dark cycle and food availability. Thus, feeding is subjected to both homeostatic and circadian regulation mechanisms that determine the amount and timing of spontaneous food intake in normal conditions. In the present study we contrasted the influence of the homeostatic versus the chronostatic mechanisms on food intake in normal conditions and in response to fasting. A group of rats was subjected to food deprivation under two different temporal schemes. A constant-length 24-h food deprivation started at different times of day resulted in an increased compensatory intake. This compensatory response showed a circadian variation that resembled the rhythm of intake in non-deprived animals. When subjected to fasting periods of increasing length (24–66 h), the amount of compensatory feeding varied according to the time of day in which food was made available, being significantly less when the fast ended in the middle of the light phase or beginning of the dark phase. These oscillatory changes did not have a correlation with variations in the level of glucose or β-hydroxybutyrate in the blood. The results suggest that the mechanism of homeostatic compensation is modulated chronostatically, presumably as part of the alternation of catabolic and anabolic states matching the daily cycles of activity.