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.     

Locusts as model organisms in which to study immunogen-induced anorectic behaviour

When injected into adult or nymphal Locusta that have been deprived of food for 2 h, immunogens such as laminarin and bacterial LPS can induce an almost immediate dose-dependent state of anorexia for at least 1 h. Such anorexia is a component of a medley of physiological and behavioural changes called collectively ‘sickness behaviour’ that occurs in a wide range of animals in response to infection or immune challenge. Sub-optimal amounts of injected laminarin allow some locusts to feed, but with a longer latency than in controls, although the length of the first meal is unaffected. The feeding behaviour of fifth instar nymphs is more sensitive to laminarin than that of adults, but both stages respond to amounts of immunogen that are lower than those required to activate the phenoloxidase cascade. Injection of adipokinetic hormone (AKH) before the period of food deprivation prevents the anorexigenic action of the laminarin in adults but not in nymphs. It is argued that the effect of the AKH may be indirect, through its lipid-mobilising action. The insecticide pymetrozine increases the latency to feed but also reduces the length of the first meal, and its anorexigenic activity is not affected by injection of AKH. The present data support the concept that laminarin-induced anorexia involves a central lack of motivation to eat, rather than a ‘stop eating’ signal. Others have shown that the mechanism of action of pymetrozine involves the serotonergic system and can be blocked by mianserin, so it is intriguing that in the present study injection of mianserin prior to that of laminarin modulates the anorexigenic effect of the immunogen. This suggests that biogenic amines are involved in the control of appetitive behaviour in locusts, as they are in vertebrates. The possible usefulness of the locust model in studying sickness-induced anorexia is discussed briefly.     

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.      

西藏飞蝗触角感器的扫描电镜观察

应用扫描电镜对西藏飞蝗Locusta migratoria tibetensis Chen触角的外部形态结构及其感器进行了观察和研究.结果表明,西藏飞蝗触角上存在5种感器即毛形感器、刺形感器、腔锥形感器、锥形感器和腔形感器.通过对各种感器的形态特点进行描述,发现西藏飞蝗群居型与散居型、蝗蝻与成虫、雌性与雄性的感器在类型...     

Mechanisms regulating salt intake in fifth-instar nymphs of Locusta migratoria

Abstract. Investigations were undertaken into the mechanisms controlling regulation of salt intake in fifth-instar nymphs of Locusta migratoria L. Initially, the pattern of feeding was recorded for 24 h in locusts fed a chemically-defined food either with or without salt. Differences were apparent from the first meal, with smaller meals being eaten more frequently on the salt-free food over the next 8h. As a result, total food intake over the first 8h was similar on the two foods. By 24 h, however, intake of the salt-free food was considerably reduced and the rate of prefeeding rejections was high. Next, following a period of pretreatment (24 or 48 h) on one of these two foods, responses were recorded to glass-fibre discs impregnated with salts, artificial foods differing in salt content, and wheat germ. Haemolymph amino acid titres were also recorded in locusts feeding ad libitum on salt-free or salt-containing food to test the 'metabolic bottleneck' hypothesis for explaining rejection of a salt-free food after a period of feeding on it. Finally, the role of learning was investigated in experiments in which distinctive plant odours and flavours were paired with foods lacking or containing salt. Results across all the experiments indicate that regulation of mineral intake in locust nymphs involves a number of mechanisms, including modulation of locomotory behaviour, innate taste responses and learning. Additionally, there was evidence of a sex difference in salt appetite.     

Amino acids in nectar enhance longevity of female Culex quinquefasciatus mosquitoes

Culex mosquitoes feed on a wide range of nectars consisting of mostly carbohydrates and amino acids, however, little is known about the utilization and effects of these different carbohydrates and their accompanying amino acids on longevity. Culex quinquefasciatus larvae were reared on low- and high-quantity food diets to produce adults that were nutritionally representative of wild-caught and laboratory-reared mosquitoes. Emerging adults reared on low- or high-quantity food diets as larvae were then provided Lantana camara nectar mimics containing mixtures of carbohydrates and amino acids to evaluate effects of nectar amino acids on longevity. Carbohydrates (with or without amino acids) were a critical component of the adult diet, and in their absence, adult mosquitoes died within 3-5 days. The nectar mimic that contained both carbohydrates and amino acids did not increase adult longevity of males originating from either poorly or well-fed larvae. However, females receiving adult diets containing both carbohydrates and amino acids lived 5% longer than females fed adult diets with only sugar.     

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.     

A Behavioral Mechanism for Incorporating an Unpalatable Food in the Diet of a Generalist Herbivore (Orthoptera: Acrididae)

We investigated mechanisms that could lead to incorporation of unpalatable foods into the diet of a generalist grasshopper, Schistocerca americana: nutritional stress, habituation, learning, and attraction to novelty. The model system involved mesquite, a palatable but inferior food, and mulberry, an unpalatable but adequate food. Nutritional stress, due to prolonged intake of the inferior food, mesquite, did not increase the acceptability of mulberry. Habituation to the deterrent compounds in mulberry and associative learning of the nutritional benefits of mulberry also did not occur. However, mulberry became more acceptable after a day of restriction to a single food type other than mulberry, and even deterrent and nutritionally worthless alternatives such as filter paper became acceptable after a day on any one food type. A tendency to feed on novel food types may be a proximate mechanism for the incorporation of relatively unpalatable, but nutritionally valuable foods into the diet. Novelty and the apparent need for diversity of foods are discussed in the context of exploratory foraging behavior by generalist herbivores.     

A correlation between macronutrient balancing and insect host-plant range: evidence from the specialist caterpillar Spodoptera exempta (Walker)

In an earlier study, we showed that the ingestive responses of the generalist caterpillar Spodoptera littoralis to foods imbalanced in their protein:carbohydrate content is similar to generalist locusts, but differs from that of specialist-feeding locusts. Here we further pursued the comparison by repeating the experiments using a closely related specialist caterpillar, Spodoptera exempta. First, caterpillars were allowed to self-compose a diet of preferred protein:carbohydrate balance by mixing between nutritionally complementary foods. Then, they were confined to one of five imbalanced foods, in which we measured the trade-off between over- and under-ingesting the two nutrients. On complementary foods, the caterpillars actively regulated their protein and carbohydrate intake. In the no-choice experiment, those fed excess-protein foods ingested small surpluses of protein compared with generalist feeders, thus showing a pattern of nutrient balancing similar to that observed in specialist locusts. Utilisation data indicated that ingested excesses and deficits were to some extent offset by differential utilisation. Evidence also showed that post-ingestive responses of the specialist S. exempta were less flexible than those observed in the generalist S. littoralis, a pattern which is again in accordance with comparisons of acridids differing in their host-plant range.     

Effect of dietary history and algal traits on feeding rate and food preference in the green sea urchin Strongylocentrotus droebachiensis

Feeding behaviour is influenced by a variety of factors, including nutritional requirements, the quality of available foods, and environmental conditions. We examined the effect of two factors, food morphology and dietary history, on the feeding rate and preference of the sea urchin Strongylocentrotus droebachiensis. Standardizing food shape and structure did not alter urchins' expected preference for the native kelp Laminaria longicruris over the invasive alga Codium fragile ssp. tomentosoides. However, when foods containing L. longicuris were shaped to mimic the algae, the C. fragile mimic was consumed more rapidly than the kelp mimic. Dietary history had no effect on single diet feeding rate. Urchins feeding on C. fragile consistently consumed twice as much (by mass) as those fed kelp, regardless of their previous diet. Despite higher feeding rates on C. fragile, urchins feeding on this alga were unable to compensate for its low energetic content and ingested less energy. Dietary history had a short-term effect on food preference, with urchins tending to prefer less familiar foods. Our findings suggest that urchins feed on C. fragile at a high rate, due to ease of handling and/or compensatory feeding, and that they do not a have strict preference hierarchy. Rather, food choice appears to reflect active maintenance of a mixed diet.     

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

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

An excitatory state generated during feeding in the locust, Chortoicetes terminifera

Various sequences of 2 μl drops of 1·0 M sucrose and water were applied directly to the mouthparts of restrained adults of the locust Chortoicetes terminifera which had been deprived of all food for 24 hr, and the number of drops consumed was recorded. Locusts fed either single drops of sucrose solution and water in alternation or single drops of sucrose solution alternated with groups of four consecutive drops of water, consumed larger total volumes than locusts fed only drops of water or locusts offered water after having been fed partially or wholly to repletion with 1·0 M sucrose. Towards the end of meals consisting of repeated cycles of one drop of sucrose solution and four drops of water, locusts consumed the one or two drops of water which followed a sucrose drop more quickly than the drop which preceded it. Furthermore, a few locusts which received this treatment terminated their meal while either a drop of sucrose solution or the water drop which immediately followed the sucrose drop was in contact with their mouthparts. The findings provide strong evidence that stimulation received during ingestion of sucrose induces, in addition to an immediate excitatory effect on the feeding system, a perseverating state of neural excitation which, for a short time afterward, increases the chance that the insect will ingest water.     

A comparison of dietary selection behaviour in larval Locusta migratoria and Spodoptera littoralis

ABSTRACT. . Final instar nymphs of the oligophagous acridid Locusta migratoria (L.) and larvae of the polyphagous noctuid Spodoptera littoralis (Boisduval) were fed for 4, 8 or 12 h, the conditioning period, on one of four artificial diets. Of these, diet PC contained 20% protein and 10% digestible carbohydrate; another, P, contained 20% protein but with the digestible carbohydrate component replaced by cellulose; a third, C, had the protein component substituted by cellulose, and the fourth, O, had both protein and digestible carbohydrate replaced. After this conditioning period, insects were given a choice of two diets, P and C, and hence an opportunity to select for the nutrients, if any, which were lacking in their previous food. Amounts eaten and selection behaviour were then recorded in detail for a total of 9 h. This paper deals with total amounts of diet eaten during the conditioning and choice periods. Spodoptera larvae were more sensitive than the locusts to being fed a nutritionally inadequate conditioning diet, and ate only small quantities of the P, C and O diets as compared with the PC diet, irrespective of the duration of conditioning. Locusts, on the other hand, when restricted to the P diet continued to eat relatively large amounts of it throughout a 12 h conditioning period. Those nymphs fed the C diet ingested large quantities (more than of the PC diet) up until 8 h, after which intake fell. When offered a choice, both species selected for the nutrients missing from the conditioning diet, even if the conditioning period had been as short as 4 h. During the first hour of choice locusts selected the P diet if they had been previously fed C and the C diet if previously fed P. Those deprived of both nutrients increased consumption of both P and C diets. Spodoptera larvae were more sensitive to prior deprivation of digestible carbohydrate than of protein. During the first hour of choice they selected the C diet if previously fed P or O but did not choose the P diet if previously fed C. In the subsequent 8 h of choice, however, a strong selection for the P diet after previous deprivation became apparent. In the locust, the selection for nutrients missing from the conditioning diet continued for the following 8 h of choice but became masked by a tendency, shown by all nymphs, to select C over P. The functional significance and possible physiological basis of all these responses is discussed.     

Role of galeal sensilla in host recognition and feeding behaviour of the Colorado potato beetle

Abstract. Close-range interactions with plants and the early stages of feeding behaviour of adult Colorado potato beetles, Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), were studied using beetles with and without various mouthpart sensilla. Representative host, potato (Solarium tuberosum) , and non-host, tomato (Lycopersicon esculentum) , leaves were treated very differently by beetles with and without their galeal or palpal sensilla. Galeal sensilla were particularly important for beetles interacting with these two plants. Compared with normal beetles, fewer beetles without galeae fed on potato leaves, and those that did feed spent more time assessing the plant and took more bites before feeding. On tomtato, more beetles without galeae fed, spent less time assessing the plant and took fewer bites before feeding. Beetles without galeae also had lower consumption rates on potato and higher consumption rates on tomato. Beetles were very reluctant to feed on tomato if the galeae were present so removal of maxillary sensilla was studied only on potato. Without the sensilla on the maxillary palpi only the time between first touch and first bite was affected (lengthened). The importance of galeal sensilla in host recognition by this species and the implications for chemosensory studies are discussed here and in a companion paper.     

Dietary glycerol and adult access to water: effects on fecundity and longevity in the almond moth

The quality of food eaten by larval insects will affect traits such as gamete production, fat reserves, muscle bulk and body size in the adult. Moreover, larvae also depend on high moisture content in the diet for survival. The almond moth (Ephestia cautella) (W.) (Lepidoptera; Pyralidae) does not feed as an adult although it continues to drink water. We tested the idea that an almond moth could compensate for a low-water diet as a larva by increasing its water intake as an adult. We reared larvae on two different food sources with different moisture regimes; standard laboratory diet with glycerol (relatively wet) and standard diet without glycerol (relatively dry). Half the adult moths from each treatment were given water to drink before their first and only mating. Our results show that wet larval diets (i.e. containing glycerol) significantly decreased fecundity (total number of eggs laid and the proportion of hatched larvae), whilst it significantly increased male and female longevity. The interaction effect of water access for adult males and females was significant, independent of the glycerol in the larval diet. Longevity in females that were not presented with water as adults was slightly higher if mated with a male that had had access to water, suggesting a mating donation of water. However, females that received water as adults showed a decreased longevity if mated with a male who had also had access to water as an adult, indicating a negative effect of water if received by both males and females. In addition, when the larval diet included glycerol, increased number of eggs laid decreased female longevity, whilst an absence of glycerol in the larval diet resulted in low female longevity that was unlinked with fecundity. Glycerol is used in many artificial insect diets and the fact that it shows a strong effect on key life-history traits (reproductive output and longevity in this species), merits careful re-examination of its effects on these important traits in other laboratory models. We also discuss the possibility that larval diet can affect female reproductive decisions.     

Nutritional Ecology of <Emphasis Type="Italic">Ateles chamek</Emphasis> in lowland Bolivia: How Macronutrient Balancing Influences Food Choices

All free-living animals must make choices regarding which foods to eat, with the choices influencing their health and fitness. An important goal in nutritional ecology is therefore to understand what governs animals’ diet selection. Despite large variation in the availability of different food items, Peruvian spider monkeys (Ateles chamek) maintain a relatively stable daily protein intake, but allow total energy intake to vary as a function of the composition of available food items. This is referred to as protein-dominated macronutrient balancing. Here we assess the influence of this nutritional strategy on daily and seasonal nutritional intakes, estimate the nutritional value of different foods, and interpret unusual food choices. We conducted continuous all-day observations of focal spider monkeys inhabiting a semideciduous forest in Bolivia. We recorded feeding events, collected foods, and analyzed their nutrient content. By using the Geometric Framework for nutrition, we show that individuals reached their daily end-point in nutrient space —balance between protein and nonprotein energy intake— by consuming nutritionally balanced foods or by alternating between nutritionally complementary foods. The macronutritionally balanced figs of Ficus boliviana were their primary staple food and therefore dominated their overall nutritional intake. Our results also demonstrate that spider monkeys consumed a diverse array of ripe fruits to overcome periods of fig scarcity rather than vice versa; they could obtain sufficient protein on a diet of pure fruit; and unripe figs constituted a nutritionally rewarding and reliable food resource. We hope that the approaches taken and the conclusions reached in this study will catalyze further inquiries into the nutritional ecology of frugivorous primates.     

Phenotypic plasticity in numbers of antennal chemoreceptors in a grasshopper: effects of food

Grasshoppers, Schistocerca americana, reared from hatching on artificial diet had fewer sensilla on the antennae in the final larval stage than insects reared on lettuce. This was true of basiconic and coeloconic sensilla (presumed olfactory) and trichoid sensilla (presumed gustatory). The degree of difference varied along the antenna and with sensillum type. Adding salicin to the diet restored the numbers of all types of sensillum to levels equal to, or approaching, those in lettuce-fed insects. The addition of some volatile compounds – carvone (monoterpene), chalcone (flavonoid), citral (monoterpene) and guaiacol (phenolic) – resulted in slight increases in number, but coumarin (phenylpropanoid) had no effect. None of the compounds, either singly or in combination, produced more sensilla than were present in plant-fed insects. Accepted: 26 January 1998     

Assimilation of carbon and nitrogen from pollen and nectar by a predaceous larva and its effects on growth and development

Abstract. 1. Predaceous insects may benefit from feeding on non‐prey foods, such as pollen, nectar, and honeydew, because they can provide nutrients that help maintain metabolism and enhance overall nutrient intake. Yet, the extent to which predaceous insects can assimilate non‐prey food and the importance of diet mixing during particular life history stages is poorly understood. In this study the relative contribution of an omnivorous diet to the growth and survivorship of a predaceous larva was tested in a hypothetical situation in which nutritionally optimal prey was not available. The study system comprised a predaceous larva (second‐ and third‐instar larvae of the green lacewing Chrysoperla carnea), nutritionally poor prey (larvae of Drosophila melanogaster), and non‐prey food (pollen suspension, a mixture of bee pollen and artificial nectar (1 M sucrose solution)). Chrysoperla carnea larvae in the mixed diet treatment were provided with both Drosophila larvae and pollen suspension, while those reared on the prey and non‐prey diet treatments received only Drosophila larvae or pollen suspension respectively. 2. The inclusion of pollen and sucrose in their diet enhanced the growth of C. carnea larvae. Second instars reared on the mixed diet developed significantly faster than their cohorts reared on the prey diet, however third instars reared on the mixed diet did not develop faster than their cohorts reared on the prey diet. Larvae reared on the mixed diet became larger adults than did those reared on either the prey or non‐prey diets. Third instars reared on the non‐prey diet completed their development while second instars in the non‐prey diet treatment failed to pupate. 3. Stable isotope analysis indicated that the larvae obtained most of their carbon (55–73%) and nitrogen (71–73%) from Drosophila but acquired only a minor amount of carbon (2–5%) and nitrogen (3–11%) from pollen. Larvae reared on the mixed and non‐prey diets acquired a relatively significant amount of carbon (23–51%) from sucrose. 4. A model, which included a novel fractionation factor to account for the isotopic effect of metamorphosis, was developed to explain the proportion of larval growth attributable to each diet item. It explained the adult δ¹³C values to within 0.2‰ and adult δ¹⁵N values to within 0.7‰ in all treatments. 5. Adults fed ¹⁵N‐labelled pollen as larvae retained the ¹⁵N signal of the pollen as adults. 6. The collective results of this study support the view that, despite their dependence on prey arthropods to obtain most of their dietary nitrogen, omnivorous lacewing larvae can enhance their growth and development by supplementing their diets with alternative non‐prey food resources. This finding is consistent with the notion that omnivory has evolved as a feeding strategy to acquire both additional nitrogen as well as trace nutrients.     

The effect of exposure to oestrous ewes on rams' sexual behaviour, plasma testosterone concentration and ability to stimulate ovulation in seasonally anoestrous ewes

Ruminants eat a variety of foods from different locations in the environment. While water, cover, social interactions, and predators are all likely to influence choice of foraging location, differences in macronutrient content among forages may also cause ruminants to forage in different locations even during a meal. We hypothesized that lambs forage at locations containing foods that complement their basal diet and meet their nutritional needs. Based on this hypothesis, we predicted that lambs (n=12) fed a basal diet low in protein and high in energy would forage where a high-protein food (Food P) was located, and that lambs (n=12) fed a basal diet low in energy and high in protein would forage where a high-energy food (Food E) was located. Food P was a ground mixture of blood meal (50%), grape pomace (30%), and alfalfa (20%) that contained 47% crude protein (CP) and 2.211 Mcal/kg digestible energy (DE). Food E was a ground mixture of cornstarch (50%), grape pomace (30%), and rolled barley (20%) that contained 6% CP and 3.07 Mcal/kg DE. Food P provided 212 g CP/Mcal DE, whereas Food E provided 20 g CP/Mcal DE. Lambs growing at a moderate rate require 179 g CP and 3.95 Mcal DE. During Trial 1, we determined if lambs foraged to correct a nutrient imbalance, and if they preferred a variety of foods (Foods P and E) to only one food at a location (Food P or E). During Trial 2, we determined if nutrient-imbalanced lambs foraged in the location with the food that corrected the imbalance when the location of the foods changed daily. During Trial 3, lambs were offered familiar foods (Foods P and E) at the location furthest - and novel foods (wheat and soybean meal) at the location nearest - the shelter of their pen. During all three trials, lambs foraged most at the location with the food that contained the highest concentration of the macronutrient lacking in their basal diet, but they always ate some of both foods. Lambs did not feed exclusively at the location with a variety of foods (P and E). Rather, they fed at the location nearest the shelter that contained the macronutrient lacking in their diet. As availability of the food with the needed macronutrient declined in one location, lambs moved to the nearest location that had food with the needed macronutrient. When food that complemented their basal diet was moved to a different location, lambs foraged in the new location. Collectively, these results show that lambs challenged by imbalances in energy or protein selected foods and foraging locations that complemented the nutrient content of their macronutrient imbalanced basal diets.