Functional implications of omnivory for dietary nutrient balance

Captive experiments have shown that many species regulate their macronutrient (i.e. protein, lipid and carbohydrate) intake by selecting complementary food types, but the relationships between foraging strategies in the wild and nutrient regulation remain poorly understood. Using the pine marten as a model species, we collated available data from the literature to investigate effects of seasonal and geographic variation in diet on dietary macronutrient balance. Our analysis showed that despite a high variety of foods comprising the diet, typical of a generalist predator, the macronutrient energy ratios of pine martens were limited to a range of 50–55% of protein, 38–42% of lipids and 5–10% of carbohydrates. This broad annual stabilisation of macronutrient ratios was achieved by using alternative animal foods to compensate for the high fluctuation of particular prey items, and sourcing non‐protein energy (carbohydrates and fats) from plant‐derived foods, particularly fruits. Macronutrient balance varied seasonally, with higher carbohydrate intake in summer–autumn, due to opportunistic fruit consumption, and higher protein intake in winter–spring. In terms of their proportional dietary carbohydrate intake the pine marten's nutritional strategy fell between that of true carnivores (e.g. the wolf) and more omnivorous feeders (e.g. the European badger). However, in terms of energy contributed by protein pine martens are equivalent to obligate carnivores such as the wolf and domesticated cat, and different to some omnivorous carnivores such as the domesticated dog and grizzly bears.     

Balancing of lipid,protein, and carbohydrate intake in a predatory beetle following hibernation,and consequences for lipid restoration

Carnivorous animals are known to balance their consumption of lipid and protein, and recent studies indicate that some mammalian carnivores also regulate their intake of carbohydrate. We investigated macronutrient balancing and lipid restoration following hibernation in the ground beetle Anchomenus dorsalis, hypothesizing that carbohydrates might be important energy sources upon hibernation when predator lipid stores are exhausted and prey are equally lean. We recorded the consumption of lipid, protein, and carbohydrate over nine days following hibernation, as the beetles foraged to refill their lipid stores. Each beetle was given the opportunity to regulate consumption from two semi-artificial foods differing in the proportion of two of the three macronutrients, while the third macronutrient was kept constant. When analyzing consumption of the three macronutrients on an energetic basis, it became apparent that the beetles regulated lipid and carbohydrate energy interchangeably and balanced the combined energy intake from the two macronutrients against protein intake. Restoration of lipid stores was independent of the availability of any specific macronutrient. However, the energetic consumption required to refill lipid stores was higher when a low proportion of lipids was ingested, suggesting that lipids were readily converted into lipid stores while there were energetic costs associated with converting carbohydrate and protein into stored lipids. Our experiment demonstrates that carbohydrates are consumed and regulated as a non-protein energy source by A. dorsalis despite an expectedly low occurrence of carbohydrates in their natural diet. Perhaps carbohydrates are in fact an overlooked supplementary energy source in the diet of carnivorous arthropods.     

Dietary protein content alters energy expenditure and composition of the mass gain in grizzly bears (Ursus arctos horribilis)

Many fruits contain high levels of available energy but very low levels of protein and other nutrients. The discrepancy between available energy and protein creates a physiological paradox for many animals consuming high-fruit diets, as they will be protein deficient if they eat to meet their minimum energy requirement. We fed young grizzly bears both high-energy pelleted and fruit diets containing from 1.6% to 15.4% protein to examine the role of diet-induced thermogenesis and fat synthesis in dealing with high-energy-low-protein diets. Digestible energy intake at mass maintenance increased 2.1 times, and composition of the gain changed from primarily lean mass to entirely fat when the protein content of the diet decreased from 15.4% to 1.6%. Daily fat gain was up to three times higher in bears fed low-protein diets ad lib., compared with bears consuming the higher-protein diet and gaining mass at the same rate. Thus, bears eating fruit can either consume other foods to increase dietary protein content and reduce energy expenditure, intake, and potentially foraging time or overeat high-fruit diets and use diet-induced thermogenesis and fat synthesis to deal with their skewed energy-to-protein ratio. These are not discrete options but a continuum that creates numerous solutions for balancing energy expenditure, intake, foraging time, fat accumulation, and ultimately fitness, depending on food availability, foraging efficiency, bear size, and body condition.     

Complementary food resources of carnivory and frugivory affect local abundance of an omnivorous carnivore

A major unresolved question for omnivorous carnivores, like most species of bears, is to what degree are populations influenced by bottom–up (food supply) or top–down (human‐caused mortality) processes. Most previous work on bear populations has focused on factors that limit survival (top–down) assuming little effect of food resource supply. When food resources are considered, most often they consider only the availability/supply of a single resource, particularly marine‐subsidized or terrestrial sources of protein (carnivory) or alternately hard or soft mast (frugivory). Little has been done to compare the importance of each of these factors for omnivorous bears or test whether complementary resources better explain individual animal and population measures such as density, vital rates, and body size. We compared landscape patterns of digestible energy (kcal) for buffaloberry (a key source of carbohydrate) and ungulate matter (a key source of protein and lipid) to local measures in grizzly bear Ursus arctos abundance at DNA hair snag sites in west‐central Alberta, Canada. We tested support for bottom–up hypotheses in either single (carnivory [meat] versus frugivory [fruit]) or complementary (additive or multiplicative) food resources, while accounting for a well‐known top–down limiting factor affecting bear survival (road density). We found support for both top–down and bottom–up factors with complementary resources (co‐limitation) supported over single resource supplies of either meat or fruit. Our study suggests that the availability of food resources that provide complementary nutrients is more important in predicting local bear abundance than single foods or nutrients (e.g. protein) or simply energy per se. This suggests a nutritionally multidimensional bottom–up limitation for a low density interior population of grizzly bears.     

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.     

The Nutritional Balancing Act of a Large Herbivore: An Experiment with Captive Moose (Alces alces L)

The nutrient balancing hypothesis proposes that, when sufficient food is available, the primary goal of animal diet selection is to obtain a nutritionally balanced diet. This hypothesis can be tested using the Geometric Framework for nutrition (GF). The GF enables researchers to study patterns of nutrient intake (e.g. macronutrients; protein, carbohydrates, fat), interactions between the different nutrients, and how an animal resolves the potential conflict between over-eating one or more nutrients and under-eating others during periods of dietary imbalance. Using the moose (Alces alces L.), a model species in the development of herbivore foraging theory, we conducted a feeding experiment guided by the GF, combining continuous observations of six captive moose with analysis of the macronutritional composition of foods. We identified the moose’s self-selected macronutrient target by allowing them to compose a diet by mixing two nutritionally complementary pellet types plus limited access to Salix browse. Such periods of free choice were intermixed with periods when they were restricted to one of the two pellet types plus Salix browse. Our observations of food intake by moose given free choice lend support to the nutrient balancing hypothesis, as the moose combined the foods in specific proportions that provided a particular ratio and amount of macronutrients. When restricted to either of two diets comprising a single pellet type, the moose i) maintained a relatively stable intake of non-protein energy while allowing protein intakes to vary with food composition, and ii) increased their intake of the food item that most closely resembled the self-selected macronutrient intake from the free choice periods, namely Salix browse. We place our results in the context of the nutritional strategy of the moose, ruminant physiology and the categorization of food quality.     

The ‘omnivorous badger dilemma’: towards an integration of nutrition with the dietary niche in wild mammals

相似文献   

The effects of nutritional imbalance on compensatory feeding for cellulose-mediated dietary dilution in a generalist caterpillar

Abstract. The interactive effects of macronutrient balance [protein (P) : carbohydrate (C) ratio] and dietary dilution by cellulose on nutritional regulation and performance were investigated in the generalist caterpillar Spodoptera littoralis (Boisduval). Caterpillars were reared through the final stadium on one of 20 foods varying factorially in macronutrient content (P + C%: 42, 33.6. 25.2 or 16.8%) and P : C ratio (5 : 1, 2 : 1, 1 : 1, 1 : 2 or 1 : 5). The animals compensate by eating more of diluted foods, but suffer reduced nutrient intake in proportion to the degree of dilution. Increase in food intake with dilution is greater on balanced than imbalanced foods and this is reflected in greater reduction of dry pupal mass with dilution in the latter. Whereas dilution results in a reduction in the amount of whichever macronutrient is in excess in the food, by contrast, the ability to compensate for the deficient macronutrient in the food is unaffected by nutrient imbalance. Excess protein intake due to nutritional imbalance (diets with high P : C ratios) results in a regulatory decrease in the efficiency of retention of ingested nitrogen relative to restricted protein intake on oppositely imbalanced foods (low P : C ratios). By contrast, decreased protein intake due to dietary dilution is associated with a non‐regulatory reduction in the efficiency of retention, irrespective of P : C ratio. Dilution is similarly associated with reduced utilization efficiency of ingested carbohydrate. The ecological implications of these results are discussed.     

Constraints on herbivory by grizzly bears

Although well known as carnivores and not capable of digesting plant fiber, grizzly bears (Ursus arctos horribilis) consume over 200 species of plants and are entirely vegetarian in some ecosystems. Even in ecosystems with abundant meat resources, green vegetation can be an important seasonal food resource. Therefore, we examined the morphological, physiological, and environmental constraints that determine the nutritional value of herbaceous vegetation to grizzly bears. Short-term, board foraging trials were used with captive grizzly bears to determine constraints on intake rate including bite size, bite rate, bear size, plant species, plant height, and plant distribution. Feeding trials were conducted to determine the effect of protein level (12-35%) and digestible dry matter intake on weight gain. Finally, maximum daily intake, daily foraging time, and weight change were measured for captive bears foraging on highly abundant and nutritious forbs and grasses during 12-day trials. Intake during short-term board trials overestimated the intake of freely foraging bears from two- to seven-fold depending on bear size. Because of their relatively larger bite sizes, smaller absolute energy requirements, and relatively larger intake capacity, smaller bears (<120 kg) made greater weight gains than very large bears on herbaceous vegetation. Smaller bears with ad libitum access to palatable, nutritious forbs gained weight at rates equal to wild bears. However, depending upon plant characteristics, bite sizes and available daily foraging time increasingly prevented large bears (>120 kg) from gaining weight on herbaceous vegetation. Both captive and wild bears select forbs over grasses at similar growth stages because forbs are generally higher in protein and more digestible than grasses. Therefore, the nutritional well-being of wild grizzly bears could be improved in areas where bears are largely herbivorous and, thus, relatively small by purposefully managing for nutritious forbs.     

Macronutrient balancing in free‐ranging populations of moose

At northern latitudes, large spatial and temporal variation in the nutritional composition of available foods poses challenges to wild herbivores trying to satisfy their nutrient requirements. Studies conducted in mostly captive settings have shown that animals from a variety of taxonomic groups deal with this challenge by adjusting the amounts and proportions of available food combinations to achieve a target nutrient balance. In this study, we used proportions‐based nutritional geometry to analyze the nutritional composition of rumen samples collected in winter from 481 moose (Alces alces) in southern Sweden and examine whether free‐ranging moose show comparable patterns of nutrient balancing. Our main hypothesis was that wild moose actively regulate their rumen nutrient composition to offset ecologically imposed variation in the nutritional composition of available foods. To test this, we assessed the macronutritional composition (protein, carbohydrates, and lipids) of rumen contents and commonly eaten foods, including supplementary feed, across populations with contrasting winter diets, spanning an area of approximately 10,000 km². Our results suggest that moose balanced the macronutrient composition of their rumen, with the rumen contents having consistently similar proportional relationship between protein and nonstructural carbohydrates, despite differences in available (and eaten) foods. Furthermore, we found that rumen macronutrient balance was tightly related to ingested levels of dietary fiber (cellulose and hemicellulose), such that the greater the fiber content, the less protein was present in the rumen compared with nonstructural carbohydrates. Our results also suggest that moose benefit from access to a greater variety of trees, shrubs, herbs, and grasses, which provides them with a larger nutritional space to maneuver within. Our findings provide novel theoretical insights into a model species for ungulate nutritional ecology, while also generating data of direct relevance to wildlife and forest management, such as silvicultural or supplementary feeding practices.     

Food limitation and starvation independently affect predator macronutrient selection

Predators are often food limited in their habitat, and some are limited by specific macronutrients (protein, lipid or carbohydrate). It is unresolved, however, to what extent and in what way food and macronutrient limitation are connected. Using a carabid beetle (Nebria brevicollis), we compared macronutrient self-selection of the animals three times: immediately after collection in the field, after being fed to satiation and nutritional balance and after a subsequent period of starvation. Both sexes were food and females lipid limited in the field; after 7–21 days of starvation both sexes increased proportional carbohydrate intake significantly. Thus, starvation created a nutrient deficit that was different from what the animals had experienced in the field. We conclude that while macronutrient limitation in nature may be influenced by hunger due to food limitation, this is not its main determinant. A nutritional imbalance of available food may override this effect.     

Functional macronutritional generalism in a large omnivore,the brown bear

We combine a recently developed framework for describing dietary generalism with compositional data analysis to examine patterns of omnivory in a large widely distributed mammal. Using the brown bear (Ursus arctos) as a model species, we collected and analyzed data from the literature to estimate the proportions of macronutrients (protein, carbohydrate, and lipid) in the diets of bear populations. Across their range, bears consumed a diversity of foods that resulted in annual population diets that varied in macronutrient proportions, suggesting a wide fundamental macronutrient niche. The variance matrix of pairwise macronutrient log‐ratios indicated that the most variable macronutrient among diets was carbohydrate, while protein and lipid were more proportional or codependent (i.e., relatively more constant log‐ratios). Populations that consumed anthropogenic foods, such agricultural crops and supplementary feed (e.g., corn), had a higher geometric mean proportion of carbohydrate, and lower proportion of protein, in annual diets. Seasonally, mean diets were lower in protein and higher in carbohydrate, during autumn compared to spring. Populations with anthropogenic subsidies, however, had higher mean proportions of carbohydrate and lower protein, across seasons compared to populations with natural diets. Proportions of macronutrients similar to those selected in experiments by captive brown bears, and which optimized primarily fat mass gain, were observed among hyperphagic prehibernation autumn diets. However, the majority of these were from populations consuming anthropogenic foods, while diets of natural populations were more variable and typically higher in protein. Some anthropogenic diets were close to the proportions selected by captive bears during summer. Our results suggest that omnivory in brown bears is a functional adaptation enabling them to occupy a diverse range of habitats and tolerate variation in the nutritional composition and availability of food resources. Furthermore, we show that populations consuming human‐sourced foods have different dietary macronutrient proportions relative to populations with natural diets.     

Testing protein leverage in lean humans: a randomised controlled experimental study

A significant contributor to the rising rates of human obesity is an increase in energy intake. The ‘protein leverage hypothesis’ proposes that a dominant appetite for protein in conjunction with a decline in the ratio of protein to fat and carbohydrate in the diet drives excess energy intake and could therefore promote the development of obesity. Our aim was to test the ‘protein leverage hypothesis’ in lean humans by disguising the macronutrient composition of foods offered to subjects under ad libitum feeding conditions. Energy intakes and hunger ratings were measured for 22 lean subjects studied over three 4-day periods of in-house dietary manipulation. Subjects were restricted to fixed menus in random order comprising 28 foods designed to be similar in palatability, availability, variety and sensory quality and providing 10%, 15% or 25% energy as protein. Nutrient and energy intake was calculated as the product of the amount of each food eaten and its composition. Lowering the percent protein of the diet from 15% to 10% resulted in higher (+12±4.5%, p = 0.02) total energy intake, predominantly from savoury-flavoured foods available between meals. This increased energy intake was not sufficient to maintain protein intake constant, indicating that protein leverage is incomplete. Urinary urea on the 10% and 15% protein diets did not differ statistically, nor did they differ from habitual values prior to the study. In contrast, increasing protein from 15% to 25% did not alter energy intake. On the fourth day of the trial, however, there was a greater increase in the hunger score between 1–2 h after the 10% protein breakfast versus the 25% protein breakfast (1.6±0.4 vs 25%: 0.5±0.3, p = 0.005). In our study population a change in the nutritional environment that dilutes dietary protein with carbohydrate and fat promotes overconsumption, enhancing the risk for potential weight gain.     

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.     

Seasonal habitat use and selection by grizzly bears in Northern British Columbia

We defined patterns of habitat use and selection by female grizzly bears (Ursus arctos) in the Besa-Prophet watershed of northern British Columbia. We fitted 13 adult females with Geographic Positioning System (GPS) radio-collars and monitored them between 2001 and 2004. We examined patterns of habitat selection by grizzly bears relative to topographical attributes and 3 potential surrogates of food availability: land-cover class, vegetation biomass or quality (as measured by the Normalized Difference Vegetation Index), and selection value for prey species themselves (moose [Alces alces], elk [Cervus elaphus], woodland caribou [Rangifer tarandus], Stone's sheep [Ovis dalli stonei]). Although vegetation biomass and quality, and selection values for prey were important in seasonal selection by some individual bears, land-cover class, elevation, aspect, and vegetation diversity most influenced patterns of habitat selection across grizzly bears, which rely on availability of plant foods and encounters with ungulate prey. Grizzly bears as a group avoided conifer stands and areas of low vegetation diversity, and selected for burned land-cover classes and high vegetation diversity across seasons. They also selected mid elevations from what was available within seasonal ranges. Quantifying relative use of different attributes helped place selection patterns within the context of the landscape. Grizzly bears used higher elevations (1,595 ± 31 m SE) in spring and lower elevations (1,436 ± 27 m) in fall; the range of average elevations used among individuals was highest (500 m) during the summer. During all seasons, grizzly bears most frequented aspects with high solar gain. Use was distributed across 10 land-cover classes and depended on season. Management and conservation actions must maintain a diverse habitat matrix distributed across a large elevational gradient to ensure persistence of grizzly bears as levels of human access increase in the northern Rocky Mountains. © 2011 The Wildlife Society.     

Dietary generalists and nutritional specialists: Feeding strategies of adult female blue monkeys (Cercopithecus mitis) in the Kakamega Forest,Kenya

Generalist primates eat many food types and shift their diet with changes in food availability. Variation in foods eaten may not, however, match variation in nutrient intake. We examined dietary variation in a generalist‐feeder, the blue monkey (Cercopithecus mitis), to see how dietary food intake related to variation in available food and nutrient intake. We used 371 all‐day focal follows from 24 adult females (three groups) in a wild rainforest population to quantify daily diet over 9 months. We measured food availability using vegetation surveys and phenology monitoring. We analyzed >700 food and fecal samples for macronutrient content. Subjects included 445 food items (species‐specific plant parts and insect morphotypes) in their diet. Variation in fruit consumption (percentage of diet and total kcal) tracked variation in availability, suggesting fruit was a preferred food type. Fruits also constituted the majority of the diet (by calories) and some fruit species were eaten more than expected based on relative availability. In contrast, few species of young leaves were eaten more than expected. Also, subjects ate fewer young leaves (based on calories consumed) when fruit or young leaves were more available, suggesting that young leaves served as fallback foods. Despite the broad range of foods in the diet, group differences in fiber digestibility, and variation that reflected food availability, subjects and groups converged on similar nutrient intakes (grand mean ± SD: 637.1 ± 104.7 kcal overall energy intake, 293.3 ± 46.9 kcal nonstructural carbohydrate, 147.8 ± 72.4 kcal lipid, 107.8 ± 12.9 kcal available protein, and 88.1 ± 17.5 kcal structural carbohydrate; N = 24 subjects). Thus, blue monkeys appear to be food composition generalists and nutrient intake specialists, using flexible feeding strategies to regulate nutrient intake. Findings highlight the importance of simultaneously examining dietary composition at both levels of foods and nutrients to understand primate feeding ecology.     

Evidence for density-dependent effects on body composition of a large omnivore in a changing Greater Yellowstone Ecosystem

Understanding the density-dependent processes that drive population demography in a changing world is critical in ecology, yet measuring performance–density relationships in long-lived mammalian species demands long-term data, limiting scientists' ability to observe such mechanisms. We tested performance–density relationships for an opportunistic omnivore, grizzly bears (Ursus arctos, Linnaeus, 1758) in the Greater Yellowstone Ecosystem, with estimates of body composition (lean body mass and percent body fat) serving as indicators of individual performance over two decades (2000–2020) during which time pronounced environmental changes have occurred. Several high-calorie foods for grizzly bears have mostly declined in recent decades (e.g., whitebark pine [Pinus albicaulis, Engelm, 1863]), while increasing human impacts from recreation, development, and long-term shifts in temperatures and precipitation are altering the ecosystem. We hypothesized that individual lean body mass declines as population density increases (H1), and that this effect would be more pronounced among growing individuals (H2). We also hypothesized that omnivory helps grizzly bears buffer energy intake from changing foods, with body fat levels being independent from population density and environmental changes (H3). Our analyses showed that individual lean body mass was negatively related to population density, particularly among growing-age females, supporting H1 and partially H2. In contrast, population density or sex had little effect on body fat levels and rate of accumulation, indicating that sufficient food resources were available on the landscape to accommodate successful use of shifting food sources, supporting H3. Our results offer important insights into ecological feedback mechanisms driving individual performances within a population undergoing demographic and ecosystem-level changes. However, synergistic effects of continued climate change and increased human impacts could lead to more extreme changes in food availability and affect observed population resilience mechanisms. Our findings underscore the importance of long-term studies in protected areas when investigating complex ecological relationships in an increasingly anthropogenic world.     

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.     

Optimizing protein intake as a foraging strategy to maximize mass gain in an omnivore

Energy maximization, time minimization, and linear programming models subject to various constraints have dominated foraging ecology ideas and methods for decades. However, animals must use very complex physiological processes and foraging decisions to ensure fitness that in many cases may not be adequately described by these approaches. An example of this problem occurs when brown bears, Ursus arctos, have access to both abundant salmon and fruit. Salmon are one of the most energy and nutrient dense foods available to bears. Fruits are often high in soluble carbohydrates, low to deficient in many required nutrients, and more difficult to efficiently exploit. Therefore, wild brown bears that fatten primarily on fruits without access to salmon are 50% smaller than salmon-feeding bears. Thus, we predicted based on a linear, energy-maximizing model without dietary interaction effects that wild brown bears with access to both abundant salmon and fruit would feed almost exclusively on salmon. However, wild adult females with or without accompanying offspring foraged three times longer per day on fruit than on salmon. Similarly, the relative dry matter intake of ad libitum apples and salmon by captive, adult brown bears averaged 76±5% fruit and 24±5% salmon. Captive brown bears consuming mixed diets with intermediate dietary protein levels had 60% lower maintenance energy costs, 37% to 139% higher efficiencies of mass gain, and 72% to 520% higher maximum rates of gain than when they consumed either salmon or fruit alone. These relationships were nonlinear functions of dietary protein content in which salmon and fruit provided complementary nutritional resources. Both wild and captive bears attempted to regulate total protein, energy, and carbohydrate intake within a multidimensional intake target that both maximized energy intake and mass gain.     

Diet and macronutrient niche of Asiatic black bear (Ursus thibetanus) in two regions of Nepal during summer and autumn

Relatively little is known about the nutritional ecology of omnivorous Asiatic black bears (Ursus thibetanus) in Nepal. We characterized the diet of black bears in two seasons (June–July, “summer”; and October–November “autumn”) and two study areas (Dhorpatan Hunting Reserve [DHR]; and Kailash Sacred Landscape [KSL]). We then conducted nutritional analysis of species consumed by black bears in each study area, in combination with nutritional estimates from the literature, to estimate the proportions of macronutrients (i.e., protein [P], lipid [L], and carbohydrate [C]) in the seasonal bear foods and diets, as well as their macronutrient niche breadth. We found that bamboo (Arundinaria spp.) had the highest relative frequency in both study areas and seasons. Ants and termites were found in DHR diets, but not KSL diets. One anthropogenic crop was found in DHR summer diets (Zea mays) and two were found in KSL summer diets (Z. mays; and Kodo millet [Paspalum scrobiculatum]). Other than insects, no animal prey was found in either diet. The proportions of macronutrients in diets (i.e., realized macronutrient niches) were relatively high in carbohydrate for both study areas and seasons: DHR_summer 24.1P:8.7L:67.2C; KSL_summer 16.7P:8.2L:75.1C; DHR_autumn 21.1P:10.5L:68.4C; KSH_autumn 19.0P:11.0L:70.0C. Macronutrient niche breadth was 3.1 × greater in the DHR than KSL during summer, and 4.0 × greater in the autumn, primarily due to the higher proportion of lipid in ants and termites relative to plant foods. Within‐study area differences in niche breadth were greater during summer than autumn; in the KSH the macronutrient breadth was 1.4 × greater in summer, while in the DHR it was 1.1 × greater in summer. Similarity in dietary macronutrient proportions despite differences in foods consumed and niche breadth are suggestive of foraging to reach a preferred macronutrient balance.