Halvor M. Halvorson

A stream insect detritivore violates common assumptions of threshold elemental ratio bioenergetics models

Ecologists increasingly use threshold elemental ratios (TERs) to explain and predict organism responses to altered resource C∶P or C∶N. TER calculations are grounded in diet-dependent growth, but growth data are limited for most taxa. Thus, TERs are derived instead from bioenergetics models that rely on simplifying assumptions, such as fixed organism C∶P and no P excretion at peak growth. We examined stoichiometric regulation of the stream insect detritivore Pycnopsyche lepida to assess bioenergetics model assumptions and compared bioenergetics TERC∶P estimates to those based on growth. We fed P. lepida maple and oak leaf diets along a dietary C∶P gradient (molar C∶P range = 950–4180) and measured consumption, growth, stoichiometric homeostasis (H), and elemental assimilation and growth efficiencies over a 5-wk period in the laboratory. Pycnopsyche lepida responses to varying resource C∶P depended on litter identity and were strongest among oak diets, on which growth peaked at diet C∶P = 1620. Pycnopsyche lepida fed oak litter exhibited flexible body C∶P during growth and in response to altered diet C∶P (non-strict homeostasis; H = 4.74), low P use efficiencies, and P excretion at peak growth. These trends violated common bioenergetics model assumptions and caused deviation of estimated TERC∶P from C∶P = 1620. Bioenergetics TERC∶P further varied among P. lepida of differing growth status on varying diet C∶P (overall TERC∶P range = 1030–9540). Our study identifies novel effects of nutrient enrichment and litter identity on detritivore stoichiometric regulation and supports growth-based approaches for future TER calculations.

Bridging Ecological Stoichiometry and Nutritional Geometry with homeostasis concepts and integrative models of organism nutrition

Summary 1.The role of nutrition in linking animals with their environment is increasingly seen as fundamental to explain ecological interactions. 2.The two currently predominant frameworks for exploring questions in nutritional ecology—Nutritional Geometry (NG) and Ecological Stoichiometry (ES)—share common features, but also differ in their goals and origins. NG originates from behavioural ecology using terrestrial insects as model organisms in tightly controlled feeding experiments, while ES originates from biogeochemistry focusing on the transfer of key elements across trophic levels, mainly in aquatic environments. 3.Here, we review the history of these two complementary frameworks, emphasizing the key concepts defining their respective aims, methodologies, and focal taxa to answer questions at different ecological scales. 4.We identify and explore homeostasis as a shared conceptual cornerstone of each framework that can be used to bridge knowledge gaps and for developing new hypotheses within nutritional ecology. 5.Expanding on the concept of homeostasis, we introduce dynamic energy budget (DEB) models as a general way to address homeostatic regulation at its fundamental level. 6.Specifically, we describe how a two-reserve DEB model can be used to track metabolic pathways of nutrients as well as elements and suggest that multi-reserve DEB models, when integrated and parameterized with NG and ES concepts, can form powerful components of agent-based models to predict how animal nutrition influences individual and trophic interactions in food webs. This article is protected by copyright. All rights reserved.

Comparing the Ecological Stoichiometry in Green and Brown Food Webs – A Review and Meta-analysis of Freshwater Food Webs

The framework of ecological stoichiometry was developed primarily within the context of “green” autotroph-based food webs. While stoichiometric principles also apply in “brown” detritus-based systems, these systems have been historically understudied and differ from green ones in several important aspects including carbon (C) quality and the nutrient [nitrogen (N) and phosphorus (P)] contents of food resources for consumers. In this paper, we review work over the last decade that has advanced the application of ecological stoichiometry from green to brown food webs, focusing on freshwater ecosystems. We first review three focal areas where green and brown food webs differ: (1) bottom–up controls by light and nutrient availability, (2) stoichiometric constraints on consumer growth and nutritional regulation, and (3) patterns in consumer-driven nutrient dynamics. Our review highlights the need for further study of how light and nutrient availability affect autotroph–heterotroph interactions on detritus and the subsequent effects on consumer feeding and growth. To complement this conceptual review, we formally quantified differences in stoichiometric principles between green and brown food webs using a meta-analysis across feeding studies of freshwater benthic invertebrates. From 257 datasets collated across 46 publications and several unpublished studies, we compared effect sizes (Pearson’s r) of resource N:C and P:C on growth, consumption, excretion, and egestion between herbivorous and detritivorous consumers. The meta-analysis revealed that both herbivore and detritivore growth are limited by resource N:C and P:C contents, but effect sizes only among detritivores were significantly above zero. Consumption effect sizes were negative among herbivores but positive for detritivores in the case of both N:C and P:C, indicating distinct compensatory feeding responses across resource stoichiometry gradients. Herbivore P excretion rates responded significantly positively to resource P:C, whereas detritivore N and P excretion did not respond; detritivore N and P egestion responded positively to resource N:C and P:C, respectively. Our meta-analysis highlights resource N and P contents as broadly limiting in brown and green benthic food webs, but indicates contrasting mechanisms of limitation owing to differing consumer regulation. We suggest that green and brown food webs share fundamental stoichiometric principles, while identifying specific differences toward applying ecological stoichiometry across ecosystems.

Quantity and quality limit detritivore growth: mechanisms revealed by ecological stoichiometry and co-limitation theory

Resource quantity and quality are fundamental bottom-up constraints on consumers. Best understood in autotroph-based systems, co-occurrence of these constraints may be common but remains poorly studied in detrital-based systems. Here, we used a laboratory growth experiment to test limitation of the detritivorous caddisfly larvae Pycnopsyche lepida across a concurrent gradient of oak litter quantity (food supply) and quality (phosphorus : carbon [P:C ratios]). Growth increased simultaneously with quantity and quality, indicating co-limitation across the resource gradients. We merged approaches of ecological stoichiometry and co-limitation theory, showing how co-limitation reflected shifts in C and P acquisition throughout homeostatic regulation. Increased growth was best explained by elevated consumption rates and improved P assimilation, which both increased with elevated quantity and quality. Notably, C assimilation efficiencies remained unchanged and achieved maximum 18% at low quantity despite pronounced C limitation. Detrital C recalcitrance and substantive post-assimilatory C losses probably set a minimum quantity threshold to achieve positive C balance. Above this threshold, greater quality enhanced larval growth probably by improving P assimilation toward P-intensive growth. We suggest this interplay of C and P acquisition contributes to detritivore co-limitation, highlighting quantity and quality as potential simultaneous bottom-up controls in detrital-based ecosystems, including under anthropogenic change like nutrient enrichment.

From Elements to Function: Toward Unifying Ecological Stoichiometry and Trait-Based Ecology

The theories developed in ecological stoichiometry are fundamentally based on traits. On the one hand, traits directly linked to cell/body stoichiometry, such as nutrient uptake and storage traits, as well as the associated trade-offs, have the potential to shape ecological interactions such as competition and predation within ecosystems. On the other hand, traits that indirectly influence and are influenced by nutritional requirements, such as cell/body size and growth rate, are tightly linked to organismal stoichiometry. Despite their physiological and ecological relevance, traits are seldom explicitly integrated in the framework of ecological stoichiometry and, currently, the major challenge is to connect ecological stoichiometry traits with functional traits considered in trait-based ecology. Here, we therefore explore and synthesize existing insights, to develop novel connections between ecological stoichiometry and trait-based ecology. By reviewing key traits and their elemental requirements and illustrating community and ecosystem consequences of variation in elemental balances of traits, we show that unifying the framework of ecological stoichiometry with trait-based ecology sheds light on the interplay between elements and functional traits. Linking elements with functions furthers our understanding of the complex connections between subcellular processes, species interactions, and ultimately ecosystem dynamics.