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ABSTRACT The role of animals in modulating nutrient cycling [hereafter, consumer-driven nutrient dynamics ( CND)] has been accepted as an important influence on both community structure and ecosystem function in aquatic systems. Yet there is great variability in the influence of CND across species and ecosystems, and the causes of this variation are not well understood. Here, we review and synthesize the mechanisms behind CND in fresh waters. We reviewed 131 articles on CND published between 1973 and 1 June 2015. The rate of new publications in CND has increased from 1.4 papers per year during 1973-2002 to 7.3 per year during 2003-2015. The majority of investigations are in North America with many concentrating on fish. More recent studies have focused on animal-mediated nutrient excretion rates relative to nutrient demand and indirect impacts (e.g. decomposition). We identified several mechanisms that influence CND across levels of biological organization. Factors affecting the stoichiometric plasticity of consumers, including body size, feeding history and ontogeny, play an important role in determining the impact of individual consumers on nutrient dynamics and underlie the stoichiometry of CND across time and space. The abiotic characteristics of an ecosystem affect the net impact of consumers on ecosystem processes by influencing consumer metabolic processes (e.g. consumption and excretion/egestion rates), non- CND supply of nutrients and ecosystem nutrient demand. Furthermore, the transformation and transport of elements by populations and communities of consumers also influences the flow of energy and nutrients across ecosystem boundaries. This review highlights that shifts in community composition or biomass of consumers and eco-evolutionary underpinnings can have strong effects on the functional role of consumers in ecosystem processes, yet these are relatively unexplored aspects of CND. Future research should evaluate the value of using species traits and abiotic conditions to predict and understand the effects of consumers on ecosystem-level nutrient dynamics across temporal and spatial scales. Moreover, new work in CND should strive to integrate knowledge from disparate fields of ecology and environmental science, such as physiology and ecosystem ecology, to develop a comprehensive and mechanistic understanding of the functional role of consumers. Comparative and experimental studies that develop testable hypotheses to challenge the current assumptions of CND, including consumer stoichiometric homeostasis, are needed to assess the significance of CND among species and across freshwater ecosystems. [ABSTRACT FROM AUTHOR]

1. Primary consumers play important roles in the cycling of nutrients in headwater streams, storing assimilated nutrients in growing tissue and recycling them through excretion. Although environmental conditions in most headwater streams and their surrounding terrestrial ecosystems vary considerably over the course of a year, relatively little is known about the effects of seasonality on consumer nutrient recycling these streams. In this study, we measured nitrogen accumulated through growth and excreted by the grazing snail Elimia clavaeformis (Pleuroceridae) over the course of 12 months in Walker Branch, identifying close connections between in-stream nitrogen processing and seasonal changes in the surrounding forest. 2. Nitrogen processing rates were positively correlated with ecosystem respiration, which was driven by leaf phenology on streamside trees. Snail nitrogen assimilation, growth and excretion were relatively high in spring before leaf emergence, low in summer when canopy shade was extensive and high again in autumn after leaf-fall. During the time that snails grazed primarily on epilithon (winter, spring and summer), growth and excretion rates followed changes in light and epilithon biomass. In autumn, when snails primarily grazed fallen leaves, leaf-associated microbes provided large subsidies of nitrogen for the snails. Nitrogen accumulation in snail biomass was greater in the 2 months following leaf-fall than at any other time of the year. 3. Snails were less important as nitrogen sinks than as sources of recycled nitrogen in Walker Branch. Over the course of the year, snails excreted approximately 12 times more nitrogen than they accumulated in biomass. Nitrogen accrued during growth in spring was subsequently lost in summer when primary production declined and snails underwent tissue loss. Catabolic losses represented >40% of the nitrogen excreted by the snails in summer. Net nitrogen growth efficiency (growth/assimilation), which varied with food availability, was only 8% for the entire year. Neither growth nor excretion was positively correlated with nitrogen concentrations in grazing substrata. 4. Snails achieved high standing crops and were significant contributors to nitrogen spiralling in Walker Branch. On an areal basis, nitrogen in snail biomass (mgN m-2) was two to five times greater than that in epilithon biomass, depending on the season. Snails assimilated and excreted up to 50% of the nitrogen initially taken up by autotrophs and leaf microbes, and they were likely to have additional effects on nitrogen spiraling through egestion and the cropping of assimilative biomass. Primary consumers like Elimia are important catalysts of nutrient movement through headwater streams, decreasing residence times and facilitating fluxes to downstream waters [ABSTRACT FROM AUTHOR]

It has recently been realized that sponges take up much of the dissolved organic matter ( DOM) available in the water of reefs. The energy derived from this DOM is suggested to be invested in renewing the sponge filter cells (choanocytes) every few hours, generating an outflow of detrital particulate organic matter ( POM) that is rapidly ingested by other invertebrates. By this DOM-to- POM recycling, sponges are proposed to fuel the food web of oligotrophic marine communities, including reefs, caves and deep-sea environments. In four species studied herein by electron microscopy, the POM found in the outgoing aquiferous canals had a complex composition, with large between-species differences. It may include choanocytes (0-52%), and also mesohyl cells, such as archeocytes (9-20%) and spherulous, and granular cells with inclusions (27-90%). Exocytosed vesicles also occurred. Surprisingly, to end up into the outgoing canals, the internal mesohyl cells squeezed between the epithelial cells (endopinacocytes) of the canal wall. Mesohyl cells were also able to transfer their inclusions to the endopinacocytes, which in turn extruded their acquired vesicle loads into the canal lumen. The unanticipated abundant participation of mesohyl cells and endopinacocytes in the production of POM appears to be an ordinary process that occurs continuously in the sponges, mostly related to elimination of digestion leftovers and excretion by-products. Therefore, POM is generated by sponges irrespective of whether the primary food source is particulate (evidence from this study) or DOM (previous literature). Altogether, these results indicate that the cellular mechanisms behind the relevant organic-matter recycling carried out by sponges are more diverse than initially anticipated. The varying ratios of choanocytes/mesohyl cells in the POM across species suggest that different sponge species may impact differently the energetics of food webs of the respective oligotrophic habitats where they dominate. [ABSTRACT FROM AUTHOR]

Nutrient recycling by fish can be an important part of nutrient cycles in both freshwater and marine ecosystems. As a result, understanding the mechanisms that influence excretion elemental ratios of fish is of great importance to a complete understanding of aquatic nutrient cycles. As fish consume a wide range of diets that differ in elemental composition, stoichiometric theory can inform predictions about dietary effects on excretion ratios., We conducted a meta-analysis to test the effects of diet elemental composition on consumption and nutrient excretion by fish. We examined the relationship between consumption rate and diet N : P across all laboratory studies and calculated effect sizes for each excretion metric to test for significant effects., Consumption rate of N, but not P, was significantly negatively affected by diet N : P. Effect sizes of diet elemental composition on consumption-specific excretion N, P and N : P in laboratory studies were all significantly different from 0, but effect size for raw excretion N : P was not significantly different from zero in laboratory or field surveys., Our results highlight the importance of having a mechanistic understanding of the drivers of consumer excretion rates and ratios. We suggest that more research is needed on how consumption and assimilation efficiency vary with N : P and in natural ecosystems in order to further understand mechanistic processes in consumer-driven nutrient recycling. [ABSTRACT FROM AUTHOR]