Geometric Framework For Nutrition Research Articles

Nutrigonometry I: Using Right-Angle Triangles to Quantify Nutritional Trade-Offs in Performance Landscapes.

AbstractAnimals regulate their food intake to maximize the expression of fitness traits but are forced to trade off the optimal expression of some fitness traits because of differences in the nutrient requirements of each trait ("nutritional trade-offs"). Nutritional trade-offs have been experimentally uncovered using the geometric framework for nutrition (GF). However, current analytical methods to measure such responses rely on either visual inspection or complex models of vector calculations applied to multidimensional performance landscapes, making these approaches subjective or conceptually difficult, computationally expensive, and, in some cases, inaccurate. Here, we present a simple trigonometric model to measure nutritional trade-offs in multidimensional landscapes (nutrigonometry) that relies on the trigonometric relationships of right-angle triangles and thus is both conceptually and computationally easier to understand and use than previous quantitative approaches. We applied nutrigonometry to a landmark GF data set for comparison of several standard statistical models to assess model performance in finding regions in the performance landscapes. This revealed that polynomial (Bayesian) regressions can be used for precise and accurate predictions of peaks and valleys in performance landscapes, irrespective of the underlying structure of the data (i.e., individual food intakes vs. fixed diet ratios). We then identified the known nutritional trade-off between life span and reproductive rate in terms of both nutrient balance and concentration for validation of the model. This showed that nutrigonometry enables a fast, reliable, and reproducible quantification of nutritional trade-offs in multidimensional performance landscapes, thereby broadening the potential for future developments in comparative research on the evolution of animal nutrition.

Nutrigonometry II: Experimental strategies to maximize nutritional information in multidimensional performance landscapes.

Animals regulate their nutrient consumption to maximize the expression of fitness traits with competing nutritional needs (“nutritional trade‐offs”). Nutritional trade‐offs have been studied using a response surface modeling approach known as the Geometric Framework for nutrition (GF). Current experimental design in GF studies does not explore the entire area of the nutritional space resulting in performance landscapes that may be incomplete. This hampers our ability to understand the properties of the performance landscape (e.g., peak shape) from which meaningful biological insights can be obtained. Here, I tested alternative experimental designs to explore the full range of the performance landscape in GF studies. I compared the performance of the standard GF design strategy with three alternatives: hexagonal, square, and random points grid strategies with respect to their accuracy in reconstructing baseline performance landscapes from a landmark GF dataset. I showed that standard GF design did not reconstruct the properties of baseline performance landscape appropriately particularly for traits that respond strongly to the interaction between nutrients. Moreover, the peak estimates in the reconstructed performance landscape using standard GF design were accurate in terms of the nutrient ratio but incomplete in terms of peak shape. All other grid designs provided more accurate reconstructions of the baseline performance landscape while also providing accurate estimates of nutrient ratio and peak shape. Thus, alternative experimental designs can maximize information from performance landscapes in GF studies, enabling reliable biological insights into nutritional trade‐offs and physiological limits within and across species.

The interaction of macronutrients and body composition among individuals with chronic spinal cord injury.

Changes in body composition and dietary intake occur following spinal cord injury (SCI). The Geometric Framework for Nutrition (GFN) is a tool that allows the examination of the complex relationships between multiple nutrition factors and health parameters within a single model. This study aimed to utilize the GFN to examine the associations between self-reported macronutrient intakes and body composition in persons with chronic SCI. Forty-eight individuals with chronic SCI were recruited. Participants completed and returned 3- or 5-day self-reported dietary recall sheets. Dietary intake of macronutrients (fats, proteins, and carbohydrates) were analysed. Anthropometric measures (circumferences), dual-energy x-ray absorptiometry (DXA), and magnetic resonance imaging (MRI) were used to assess whlole-body composition. Associations between all circumference measures and carbohydrates were observed. Among MRI measures, only significant associations between subcutaneous adipose tissue and protein x carbohydrate as well as carbohydrates alone were identified. Carbohydrates were negatively associated with several measures of fat mass as measured by DXA. Overall, carbohydrates appear to play an important role in body composition among individuals with SCI. Higher carbohydrate intake was associated with lower fat mass. Additional research is needed to determine how carbohydrate intake influences body composition and cardiometabolic health after SCI.

Sex-Specific Effect of the Dietary Protein to Carbohydrate Ratio on Personality in the Dubia Cockroach.

Simple SummaryAnimal personality is modulated by genetic and environmental factors. To explore the modulatory effect of nutrition on personality, we investigated whether diets varying in their relative content of proteins and carbohydrates might modulate the behavior of the Dubia cockroach. Over a period of eight weeks, we fed adult cockroaches, both males and females, five different diets, and we measured diet consumption, survival, and personality traits by recording their exploratory and mobility behaviors. After eight weeks, females gained more body mass and had higher survival than males. We found that females preferred carbohydrate-rich diets and avoided ingesting too many proteins by consuming less food on high-protein diets. The diet had no effect on their personality. However, males showed a bolder personality when fed with high-protein diets while consuming the same amount of food, regardless of the protein content in the diet. These sex differences could be beneficial for the species in stressful nutritional environments, allowing males to discover new food resources while ovoviviparous females could spend more time protected in shelters.Animal personality, defined by behavioral variations among individuals consistent over contexts or time, is shaped by genetic and environmental factors. Among these factors, nutrition can play an important role. The Geometric Framework of Nutrition has promoted a better understanding of the role of the macronutrient proportion in animal development, survival, reproduction, and behavior, and can help to disentangle its modulatory effect on animal personality. In this study, we investigated the effects of protein to carbohydrate (P:C) ratio in the personality of the cockroach Blaptica dubia. Newly emerged adults were fed over a period of eight weeks on five different diets varying in their P:C ratio and their diet consumption, mass variation, survival, exploratory behavior, and mobility were assessed. We found that females, unlike males, were able to regulate their nutrient intake and preferred carbohydrate-rich diets. Females also gained more body mass and lived longer compared to males. In addition, their behavior and mobility were not affected by the diet. In males, however, high-protein diets induced a bolder personality. We suggest that the sex-specific effects observed on both survival and behavior are related to the nutrient intake regulation capacity and might improve the species’ fitness in adverse nutritional conditions.

Intralocus sexual conflict over optimal nutrient intake and the evolution of sex differences in life span and reproduction

Abstract Despite widespread variation in life span across species, three clear patterns exist: sex differences in life span are ubiquitous, life span is commonly traded against reproduction, and nutrition has a major influence on these traits and how they trade‐off. One process that potentially unites these patterns is intralocus sexual conflict over the optimal intake of nutrients for life span and reproduction. If nutrient intake has sex‐specific effects on life span and reproduction but nutrient choice is genetically linked across the sexes, intralocus sexual conflict will occur and may prevent one or both sexes from feeding to their nutritional optima. Here we determine the potential for this process to operate in the decorated cricket Gryllodes sigillatus. Using the Geometric Framework for Nutrition, we restrict male and female crickets to diets varying in the ratio of protein to carbohydrates and total nutrient content to quantify the effects on life span and daily reproductive effort in the sexes. We then use inbred lines to estimate the quantitative genetic basis of nutrient choice in males and females. We combine the nutrient effects and genetic estimates to predict the magnitude of evolutionary constraint for these traits in each sex. Finally, we present male and female crickets with a much broader range of diet pairs to determine how the sexes actively regulate their intake of nutrients. We show that protein and carbohydrate intake have contrasting effects on life span and reproduction in the sexes and that there are strong positive intersexual genetic correlations for the intake of these nutrients under dietary choice. This is predicted to accelerate the evolutionary response of nutrient intake in males but constrain it in females, suggesting they are losing the conflict. Supporting this view, males and females regulate nutrient intake to a common nutrient ratio that was not perfectly optimal for life span or reproduction in either sex, especially in females. Our findings show that intralocus sexual conflict over the optimal intake of nutrients is likely to be an important process generating sex differences in life span and reproduction and may help explain why females age faster and live shorter than males in G. sigillatus. A free Plain Language Summary can be found within the Supporting Information of this article.

Behavioural and physiological regulation of protein and carbohydrates in mealworm larvae: A geometric analysis

Protein-carbohydrate regulation in the larvae of the mealworm beetle (Tenebrio molitor L.) was analyzed using the Geometric Framework for nutrition. In this study, the ingestive and post-ingestive responses were measured from T. molitor larvae that were subjected to choice and no-choice experiments. In the choice experiment, T. moitor larvae were simultaneously presented with one of two protein-biased foods (p35:c7 or p28:c5.6) and one of two carbohydrate-biased foods (p7:c35 or p5.6:c28). T. molitor larvae selected protein and carbohydrate in a ratio close to 1:1 over the first 15 days since the start of the experiment (days 0–15), but exhibited preference for carbohydrate-biased food over the next 15 days. The average protein:carbohydrate ratio selected over days 0–30 was 1:1.24. In the no-choice experiment, T. molitor larvae were restricted to one of seven foods with different protein and carbohydrate content (p0:c42, p7:c35, p14:c28, p21:c21, p28:c14, p35:c7, or p42:c0). On the p0:c42 food, consumption was greatly suppressed and no larvae completed their development. Across a range of these foods except p0:c42, T. molitor larvae consistently over-ate the surplus nutrient in the foods and showed a pattern of nutrient balancing similar to that previously described for other nutritional generalists. Despite having consumed substantially different amounts and ratios of macronutrients as larvae, T. molitor pupae in the no-choice food treatments had similar body nutrient composition, suggesting the presence of strong homeostatic regulation for body nutrient growth. Larval survivorship was significantly lower on two extremely imbalanced foods (p7:c35 and p42:0) than on more balanced foods. T. molitor larvae reared on p7:c35 suffered reduced biomass growth and delayed development compared with those on foods with higher protein content.

The Geometric Framework for Nutrition: New Insights Into How What We Eat Affects How We Age

The geometric framework for nutrition (GFN) is an approach to understanding the effect of nutrition considering multiple nutrients simultaneously. Originally developed in experimental studies of insects to model how nutritional needs evolve depending on ecological context, and since extended to many taxa including non-human primates in the wild, the technique is increasingly applied to understand human health and aging. Here, we invite four varied talks showcasing the flexibility and potential of this approach from the basic biology of aging to observational human studies and clinical trials. D. Raubenheimer will give an overview of the method, its history, and its applications in aging and human health. D. Wahl will present results showing how GFN can help develop diets that recapitulate caloric restriction and its effects on brain aging. S. Das will show how GFN can be used to improve the feasibility of caloric restriction in humans without compromising its effects. Finally, A. Cohen will present results showing how GFN can be deployed in an epidemiological context and used to characterize complex interactions among large numbers of nutrients in determining health. Together, these results show that a simplistic conception of nutrition as calories is far from sufficient to understand its effects on health and aging. Evolution has shaped the nutritional needs of each species for its environment, with appropriate levels of flexibility. GFN provides an approach to capture the relevant nuance, with the results presented at this symposium but scratching the surface. Nutrition Interest Group Sponsored Symposium.

Comparing the Effects of Low-Protein and High-Carbohydrate Diets and Caloric Restriction on Brain Aging in Mice

The Geometric Framework for Nutrition (GFN) has revealed that ad-libitum low-protein, high-carbohydrate (LPHC) diets improve cardiometabolic health and extend lifespan in rodents, but it is not known whether these diets are also beneficial for brain health. Here, we utilized previous results from GFN studies and compared hippocampus biology and memory in mice subjected to 20% calorie restriction (CR) or provided ad-libitum access to several LPHC diets. RNA expression in the hippocampus of 15-month-old mice were similar between mice fed CR and LPHC diets. Nutrient-sensing proteins, including SIRT1, MTOR, and PGC1-alpha, were also influenced by diet; however, the effects varied by sex. CR and LPHC diets were associated with increased dendritic spines in dentate gyrus neurons. Mice fed CR and LPHC diets had modest improvements in the Barnes maze spatial recognition memory paradigm and novel object recognition test. LPHC diets recapitulate some of the benefits of CR on brain aging. Part of a symposium sponsored by the Nutrition Interest Group.

Nutrient Intake Patterns Predict Homeostatic Dysregulation in an Older Quebec Population

The geometric framework for nutrition has largely been applied to macronutrients in experimental settings. Here, we utilize the framework to examine both macro and micronutrient intake patterns in observational human data. We used nutritional intake patterns (3x 24h recall per visit) of 1754 older Quebecers from the NuAge cohort to predict multi-system homeostatic dysregulation scores calculated from 30 biomarkers. Intermediate intake of both macro- and micronutrients was generally associated with lower dysregulation scores (i.e., better health). Furthermore, there were often nutrient-nutrient interactions, such that the optimal level of one nutrient depends on the intake level of others. However, higher protein intake was generally associated with better health, and results varied substantially across different dysregulation systems. Accordingly, even though nutrition does have important effects on health trajectories during aging, it will be challenging to arrive at population-level recommendations to fine-tune nutrient intake patterns to optimize health beyond “everything in moderation.” Part of a symposium sponsored by the Nutrition Interest Group.

Diet Composition, Adherence to Calorie Restriction, and Cardiometabolic Disease Risk Modification

Calorie restriction (CR) is a promising strategy to attenuate age-related disease risk. Higher protein diets enhance satiety but may also impair metabolic health and accelerate aging. The effect of higher protein intake on adherence to CR and cardiometabolic markers of healthspan remains unknown. We used the Geometric Framework for Nutrition to examine the association between diet composition and 1) CR adherence; and 2) cardiometabolic risk factors during a 2-year intervention. The CR group consumed higher percentage energy from protein and lower fat at 12 months compared to baseline (logit % protein=0.1; 95% CI=0.05, 0.15; logit % fat= -0.12; CI=-0.29, -0.18). Higher protein intake over the 2-year intervention was associated with higher adherence to CR. No effect of diet composition on cardiometabolic risk factors was observed. These findings suggest that dietary protein plays a critical role in adherence to CR with no adverse effects on cardiometabolic markers of healthspan. Part of a symposium sponsored by the Nutrition Interest Group.

Diet modulates the relationship between immune gene expression and functional immune responses

Nutrition is vital to health and the availability of resources has long been acknowledged as a key factor in the ability to fight off parasites, as investing in the immune system is costly. Resources have typically been considered as something of a “black box”, with the quantity of available food being used as a proxy for resource limitation. However, food is a complex mixture of macro- and micronutrients, the precise balance of which determines an animal's fitness. Here we use a state-space modelling approach, the Geometric Framework for Nutrition (GFN), to assess for the first time, how the balance and amount of nutrients affects an animal's ability to mount an immune response to a pathogenic infection.Spodoptera littoralis caterpillars were assigned to one of 20 diets that varied in the ratio of macronutrients (protein and carbohydrate) and their calorie content to cover a large region of nutrient space. Caterpillars were then handled or injected with either live or dead Xenorhabdus nematophila bacterial cells. The expression of nine genes (5 immune, 4 non-immune) was measured 20 h post immune challenge. For two of the immune genes (PPO and Lysozyme) we also measured the relevant functional immune response in the hemolymph. Gene expression and functional immune responses were then mapped against nutritional intake.The expression of all immune genes was up-regulated by injection with dead bacteria, but only those in the IMD pathway (Moricin and Relish) were substantially up-regulated by both dead and live bacterial challenge. Functional immune responses increased with the protein content of the diet but the expression of immune genes was much less predictable.Our results indicate that diet does play an important role in the ability of an animal to mount an adequate immune response, with the availability of protein being the most important predictor of the functional (physiological) immune response. Importantly, however, immune gene expression responds quite differently to functional immunity and we would caution against using gene expression as a proxy for immune investment, as it is unlikely to be reliable indicator of the immune response, except under specific dietary conditions.

Macronutrient intakes and the lifespan-fecundity trade-off: a geometric framework agent-based model.

Lifespan and fecundity, the main components in evolutionary fitness, are both strongly affected by nutritional state. Geometric framework of nutrition (GFN) experiments has shown that lifespan and fecundity are separated in nutrient space leading to a functional trade-off between the two traits. Here we develop a spatially explicit agent-based model (ABM) using the GFN to explore how ecological factors may cause selection on macronutrient appetites to optimally balance these life-history traits. We show that increasing the risk of extrinsic mortality favours intake of a mixture of nutrients that is associated with maximal fecundity at the expense of reduced longevity and that this result is robust across spatial and nutritional environments. These model behaviours are consistent with what has been observed in studies that quantify changes in life history in response to environmental manipulations. Previous GFN-derived ABMs have treated fitness as a single value. This is the first such model to instead decompose fitness into its primary component traits, longevity and fecundity, allowing evolutionary fitness to be an emergent property of the two. Our model demonstrates that selection on macronutrient appetites may affect life-history trade-offs and makes predictions that can be directly tested in artificial selection experiments.

The Geometry of Nutrient Space-Based Life-History Trade-Offs: Sex-Specific Effects of Macronutrient Intake on the Trade-Off between Encapsulation Ability and Reproductive Effort in Decorated Crickets.

Life-history theory assumes that traits compete for limited resources, resulting in trade-offs. The most commonly manipulated resource in empirical studies is the quantity or quality of diet. Recent studies using the geometric framework for nutrition, however, suggest that trade-offs are often regulated by the intake of specific nutrients, but a formal approach to identify and quantify the strength of such trade-offs is lacking. We posit that trade-offs occur whenever life-history traits are maximized in different regions of nutrient space, as evidenced by nonoverlapping 95% confidence regions of the global maximum for each trait and large angles (θ) between linear nutritional vectors and Euclidean distances (d) between global maxima. We then examined the effects of protein and carbohydrate intake on the trade-off between reproduction and aspects of immune function in male and female Gryllodes sigillatus. Female encapsulation ability and egg production increased with the intake of both nutrients, whereas male encapsulation ability increased with protein intake but calling effort increased with carbohydrate intake. The trade-offs between traits was therefore larger in males than in females, as demonstrated by significant negative correlations between the traits in males, nonoverlapping 95% confidence regions, and larger estimates of θ and d. Under dietary choice, the sexes had similar regulated intakes, but neither optimally regulated nutrient intake for maximal trait expression. We highlight the fact that greater consideration of specific nutrient intake is needed when examining nutrient space-based trade-offs.

The Geometric Framework for Nutrition as a tool in precision medicine.

Fundamental questions in nutrition include, “What constitutes a nutritionally balanced diet?”, “What are the consequences of failing to achieve diet balance?”, and “How does diet balance change across the lifecourse and with individual circumstances?”. Answering these questions requires coming to grips with the multidimensionality and dynamic nature of nutritional requirements, foods and diets, and the complex relationships between nutrition and health, while at the same time avoiding becoming overwhelmed by complexity. Here we illustrate the use of an integrating framework for taming the complexity of nutrition, the Geometric Framework for Nutrition (GFN), and show how this might be used to untap the full potential for nutrition to provide targeted primary interventions and treatments for the chronic diseases of aging. We first briefly introduce the concepts behind GFN, then provide an example of how GFN has been used to relate nutrition to various behavioural, physiological and health outcomes in a large mouse experiment, and end by suggesting a translational pathway to human health.

Intergenerational effects of nutrition on immunity: a systematic review and meta-analysis.

Diet and immunity are both highly complex processes through which organisms interact with their environment and adapt to variable conditions. Parents that are able to transmit information to their offspring about prevailing environmental conditions have a selective advantage by 'priming' the physiology of their offspring. We used a meta-analytic approach to test the effect of parental diet on offspring immune responses. Using the geometric framework for nutrition (a method for analysing diet compositions wherein food nutrient components are expressed as axes in a Cartesian coordinate space) to define dietary manipulations in terms of their energy and macronutrient compositions, we compiled the results of 226 experiments from 38 published papers on the intergenerational effects of diet on immunity, across a range of study species and immunological responses. We observed intergenerational impacts of parental nutrition on a number of offspring immunological processes, including expression of pro-inflammatory biomarkers as well as decreases in anti-inflammatory markers in response to certain parental diets. For example, across our data set as a whole (encompassing several types of dietary manipulation), dietary stress in parents was seen to significantly increase pro-inflammatory cytokine levels measured in offspring (overall d=0.575). All studies included in our analysis were from experiments in which the offspring were raised on a normal or control diet, so our findings suggest that a nutrition-dependent immune state can be inherited, and that this immune state is maintained in the short term, despite offspring returning to an 'optimal' diet. We demonstrate how the geometric framework for nutrition can be used to disentangle the role that different forms of dietary manipulation can have on intergenerational immunity. For example, offspring B-cell responses were significantly decreased when parents were raised on a range of different diets. Similarly, our approach allowed us to show that a parental diet elevated in protein (regardless of energy composition and relative to a control diet) can increase expression of inflammatory markers while decreasing B-cell-associated markers. By conducting a systematic review of the literature, we have identified important gaps that impair our understanding of the intergenerational effects of diet, such as a paucity of experimental studies involving increased protein and decreased energy, and a lack of studies directed at the whole-organism consequences of these processes, such as immune resilience to infection. The results of our analyses inform our understanding of the effects of diet on physiological state across diverse biological fields, including biomedical sciences, maintenance of agricultural breed stock and conservation breeding programs, among others.

Woodstoich III: Integrating tools of nutritional geometry and ecological stoichiometry to advance nutrient budgeting and the prediction of consumer‐driven nutrient recycling

Within the last two decades, ecological stoichiometry (ES) and nutritional geometry (NG, also known as geometric framework for nutrition) have delivered novel insights into core questions of nutritional ecology. These two nutritionally explicit frameworks differ in the ‘nutrient currency’ used and the focus of their past research; behavioural feeding strategies in NG, mainly investigating terrestrial organisms, and trophic ecology in ES, mainly in aquatic settings. However, both NG and ES have developed in explaining patterns across various scales of biological organization. Integrating specific tools of these frameworks could advance the field of nutritional ecology by unifying theoretical and empirical approaches from the organismal to ecosystem level processes. Toward this integration, we identified 1) nutrient/element budgets as a shared concept of both frameworks that encompass nutrient intake, retention, and release, 2) response surface plots of NG as powerful tools to illustrate processes at the organismal level and 3) the concept of consumer‐driven nutrient recycling (CNR) of ES as a useful tool bridging organism and ecosystem scales. We applied response surface plots to element budget data from an ES study to show how this approach can deliver new insights at the organismal level, e.g. by showing the interplay between egestion and excretion depending simultaneously on the consumed amount of carbon and phosphorus based on variation across individuals. By integrating concepts of ES and NG to model microbial uptake and mineralization of nitrogenous wastes reported in a NG study, we also demonstrate that considering biochemically explicit mineralization rates of organic wastes can improve predictions of CNR by reducing over‐ or underestimation of mineralization depending on the quality of the consumer's diet. Our presented tools and approaches can help to bridge the organismal and ecosystem level, advancing the predictive power of studies in nutritional ecology at multiple ecological scales.

Sucrose Sensitivity of Honey Bees Is Differently Affected by Dietary Protein and a Neonicotinoid Pesticide.

Over a decade, declines in honey bee colonies have raised worldwide concerns. Several potentially contributing factors have been investigated, e.g. parasites, diseases, and pesticides. Neonicotinoid pesticides have received much attention due to their intensive use in crop protection, and their adverse effects on many levels of honey bee physiology led the European Union to ban these compounds. Due to their neuronal target, a receptor expressed throughout the insect nervous system, studies have focused mainly on neuroscience and behaviour. Through the Geometric Framework of nutrition, we investigated effects of the neonicotinoid thiamethoxam on survival, food consumption and sucrose sensitivity of honey bees (Apis mellifera). Thiamethoxam did not affect protein and carbohydrate intake, but decreased responses to high concentrations of sucrose. Interestingly, when bees ate fixed unbalanced diets, dietary protein facilitated better sucrose detection. Both thiamethoxam and dietary protein influenced survival. These findings suggest that, in the presence of a pesticide and unbalanced food, honey bee health may be severely challenged. Consequences for foraging efficiency and colony activity, cornerstones of honey bee health, are also discussed.

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.

Nutrient balancing of the adult worker bumblebee (Bombus terrestris) depends on the dietary source of essential amino acids.

ABSTRACTAnimals carefully regulate the amount of protein that they consume. The quantity of individual essential amino acids (EAAs) obtained from dietary protein depends on the protein source, but how the proportion of EAAs in the diet affects nutrient balancing has rarely been studied. Recent research using the Geometric Framework for Nutrition has revealed that forager honeybees who receive much of their dietary EAAs from floral nectar and not from solid protein have relatively low requirements for dietary EAAs. Here, we examined the nutritional requirements for protein and carbohydrates of foragers of the buff-tailed bumblebee Bombus terrestris. By using protein (sodium caseinate) or an equimolar mixture of the 10 EAAs, we found that the intake target (nutritional optimum) of adult workers depended on the source and proportion of dietary EAAs. When bees consumed caseinate-containing diets in a range of ratios between 1:250 and 1:25 (protein to carbohydrate), they achieved an intake target (IT) of 1:149 (w/w). In contrast to those fed protein, bees fed the EAA diets had an IT more biased towards carbohydrates (1:560 w/w) but also had a greater risk of death than those fed caseinate. We also tested how the dietary source of EAAs affected free AAs in bee haemolymph. Bees fed diets near their IT had similar haemolymph AA profiles, whereas bees fed diets high in caseinate had elevated levels of leucine, threonine, valine and alanine in the haemolymph. We found that like honeybees, bumblebee workers prioritize carbohydrate intake and have a relatively low requirement for protein. The dietary source of EAAs influenced both the ratio of protein/EAA to carbohydrate and the overall amount of carbohydrate eaten. Our data support the idea that EAAs and carbohydrates in haemolymph are important determinants of nutritional state in insects.

Modelling nutrition across organizational levels: From individuals to superorganisms

The Geometric Framework for nutrition has been increasingly used to describe how individual animals regulate their intake of multiple nutrients to maintain target physiological states maximizing growth and reproduction. However, only a few studies have considered the potential influences of the social context in which these nutritional decisions are made. Social insects, for instance, have evolved extreme levels of nutritional interdependence in which food collection, processing, storage and disposal are performed by different individuals with different nutritional needs. These social interactions considerably complicate nutrition and raise the question of how nutrient regulation is achieved at multiple organizational levels, by individuals and groups. Here, we explore the connections between individual- and collective-level nutrition by developing a modelling framework integrating concepts of nutritional geometry into individual-based models. Using this approach, we investigate how simple nutritional interactions between individuals can mediate a range of emergent collective-level phenomena in social arthropods (insects and spiders) and provide examples of novel and empirically testable predictions. We discuss how our approach could be expanded to a wider range of species and social systems.