Consumer‐driven nutrient dynamics in freshwater ecosystems: from individuals to ecosystems

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.     

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.     

Size-fractionated productivity and nutrient dynamics of phytoplankton in subtropical coastal environments

It is now well established that the size distribution of phytoplankton plays an important role in primary production processes and nutrient dynamics of coastal environment. In situ observations showed that nanophytoplankton (3–20 μm) contributed 72.08% and58.18% of phytoplankton biomass and 58.32% and 41.14% of primary productivity to Xiamen Western Waters and the northern Taiwan Strait, respectively; picophytoplankton (0.2–3 μm) dominated the biomass (64.70%) and productivity (66.09%) in the southern Taiwan Strait. Furthermore, nanophytoplankton accounted for 75% of phosphate uptake with the highest rate constant (8.3×10^-5 s^-1) and uptake rate in unit water volume (5.4×10^-5 mmol dm^-3s^-1); picophytoplankton had the highest uptake rate in unit biomass (5.4×10^-5 mmol mg^-1s^-1) and photosynthetic index (3.8 mgC mgChl a^-1h^-1). All the results highlighted the remarkable characteristics of small size ranged (0.2–20 μm) phytoplankton in subtropical coastal environments: main contributor to phytoplankton biomass and production, high efficiency on organic carbon production and nutrient recycling. The far reaching environmental and ecological implications were discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

不同生长阶段杉木人工林土壤C∶N∶P化学计量特征与养分动态

在福建省白砂国有林场选取幼龄林(5年)、中幼龄林(8年)、近熟林(21年)、成熟林(27年)和过熟林(40年)5个生长阶段的杉木人工林,测定不同土层(0～10、10～20、20～40 cm)土壤总碳(TC)、全氮(TN)、全磷(TP)、全钾(TK)、全钙(Ca)、全镁(Mg)含量以及C∶N∶P化学计量比,探究杉木人工林土壤碳氮磷(C∶N∶P)化学计量特征与养分随林龄的变化规律。结果表明: 随着林分发育,TC、TN未发生显著变化,土壤C∶N保持不变。随着林分发育,0～20 cm土层土壤TP含量呈增加-降低-增加的变化趋势,其中在杉木成熟林达到最低,C∶P和N∶P最大,而20～40 cm土层土壤TP在各个林龄之间无显著变化。Ca、Mg含量在所有土层均在杉木成熟林达到最低。土壤TC与C∶P、N∶P、C∶N均呈显著正相关,TP与C∶P、N∶P呈显著负相关,土壤TP含量是调控土壤C∶P和N∶P的关键因子。杉木人工林发育到成熟期受到P的限制,为保证人工林正常发育,可在杉木速生阶段施加P肥,促进养分良性循环。适当提高杉木林的轮伐期可能会有利于土壤养分的恢复与保持。     

Fluctuations in water level and the dynamics of zooplankton: a data‐driven modelling approach

1. The zooplankton in Lake Kinneret (Israel) have undergone large fluctuations in recent decades, which have been linked to both biotic and abiotic processes. 2. By applying a data‐driven modelling approach to a long‐term database, and focusing on key abiotic (lake‐level change) and biotic (prey abundance) variables, we attempted to identify the possible factors impacting the lake’s zooplankton community. 3. We hypothesised that changes in the predatory zooplankton (adult cyclopoids) assemblage are driven by changes in lake level during years of large changes. We further postulated that lake‐level changes would have a similar impact on the herbivorous zooplankton (cladocerans and cyclopoid copepodites) but an opposite effect on the microzooplankton. In the years of moderate changes to lake level, however, the abundance of predatory zooplankton would determine the size of the herbivore and microzooplankton populations rather than their food sources, that is, top‐down control. 4. The resulting decision trees supported the hypotheses stressing the importance of the annual rate of lake‐level change in shaping the zooplankton community in the lake. In addition, and in contrast to expectations, bottom‐up processes seem to play a role in determining zooplankton abundance.     

Consumer‐resource stoichiometry as a predictor of trophic discrimination (Δ13C, Δ15N) in aquatic invertebrates

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Plant sizes mediate mowing‐induced changes in nutrient stoichiometry and allocation of a perennial grass in semi‐arid grassland

While mowing‐induced changes in plant traits and their effects on ecosystem functioning in semi‐arid grassland are well studied, the relations between plant size and nutrient strategies are largely unknown. Mowing may drive the shifts of plant nutrient limitation and allocation. Here, we evaluated the changes in nutrient stoichiometry and allocation with variations in sizes of Leymus chinensis, the dominant plant species in Inner Mongolia grassland, to various mowing frequencies in a 17‐yr controlled experiment. Affected by mowing, the concentrations of nitrogen (N), phosphorus (P), and carbon (C) in leaves and stems were significantly increased, negatively correlating with plant sizes. Moreover, we found significant trade‐offs between the concentrations and accumulation of N, P, and C in plant tissues. The N:P ratios of L. chinensis aboveground biomass, linearly correlating with plant size, significantly decreased with increased mowing frequencies. The ratios of C:N and C:P of L. chinensis individuals were positively correlated with plant size, showing an exponential pattern. With increased mowing frequencies, L. chinensis size was correlated with the allocation ratios of leaves to stems of N, P, and C by the tendencies of negative parabola, positive, and negative linear. The results of structure equation modeling showed that the N, P, and C allocations were co‐regulated by biomass allocation and nutrient concentration ratios of leaves to stems. In summary, we found a significant decoupling effect between plant traits and nutrient strategies along mowing frequencies. Our results reveal a mechanism for how long‐term mowing‐induced changes in concentrations, accumulations, ecological stoichiometry, and allocations of key elements are mediated by the variations in plant sizes of perennial rhizome grass.     

Animal pee in the sea: consumer‐mediated nutrient dynamics in the world's changing oceans

Humans have drastically altered the abundance of animals in marine ecosystems via exploitation. Reduced abundance can destabilize food webs, leading to cascading indirect effects that dramatically reorganize community structure and shift ecosystem function. However, the additional implications of these top‐down changes for biogeochemical cycles via consumer‐mediated nutrient dynamics (CND) are often overlooked in marine systems, particularly in coastal areas. Here, we review research that underscores the importance of this bottom‐up control at local, regional, and global scales in coastal marine ecosystems, and the potential implications of anthropogenic change to fundamentally alter these processes. We focus attention on the two primary ways consumers affect nutrient dynamics, with emphasis on implications for the nutrient capacity of ecosystems: (1) the storage and retention of nutrients in biomass, and (2) the supply of nutrients via excretion and egestion. Nutrient storage in consumer biomass may be especially important in many marine ecosystems because consumers, as opposed to producers, often dominate organismal biomass. As for nutrient supply, we emphasize how consumers enhance primary production through both press and pulse dynamics. Looking forward, we explore the importance of CDN for improving theory (e.g., ecological stoichiometry, metabolic theory, and biodiversity–ecosystem function relationships), all in the context of global environmental change. Increasing research focus on CND will likely transform our perspectives on how consumers affect the functioning of marine ecosystems.     

Ecological stoichiometry and nutrient partitioning in two insect herbivores responsible for large‐scale forest disturbance in the Fennoscandian subarctic

1. Outbreaks of herbivorous insects can have large impacts on regional soil carbon (C) storage and nutrient cycling. In northernmost Europe, population outbreaks of several geometrid moth species regularly cause large‐scale defoliation in subarctic birch forests. An improved understanding is required of how leaf C and nutrients are processed after ingestion by herbivores and what this means for the quantity and quality of different materials produced (frass, bodies). 2. In this study, larvae of two geometrid species responsible for major outbreaks (Epirrita autumnata and Operophtera brumata) were raised on exclusive diets of Betula pubescens var. czerepanovii (N. I. Orlova) Hämet Ahti and two other abundant understorey species (Betula nana, Vaccinium myrtillus). The quantities of C, nitrogen (N) and phosphorus (P) ingested and allocated to frass, bodies and (in the case of C) respired were recorded. 3. Overall, 23%, 70% and 48% of ingested C, N and P were allocated to bodies, respectively, rather than frass and (in the case of C) respiration. Operophtera brumata consistently maintained more constant body stoichiometric ratios of C, N and P than did E. autumnata, across the wide variation in physico‐chemical properties of plant diet supplied. 4. These observed differences and similarities on C and nutrient processing may improve researchers' ability to predict the amount and stoichiometry of frass and bodies generated after geometrid outbreaks.     

Leaf litter stoichiometry affects decomposition rates and nutrient dynamics in tropical forests under restoration in Costa Rica

Active restoration strategies increase the production of leaf litter in tropical forests, but little is known about their effect on litter decomposition and subsequent nutrient release. We quantified changes in leaf litter stoichiometry during decomposition in former pasture sites under contrasting restoration strategies (natural regeneration, applied nucleation/islands tree planting and plantation), as well as in nearby primary forest. Litterbags were employed to evaluate decomposition. We used a leaf mixture of either the four planted tree species in the plantation and island treatments or the nearby primary forest and compared them under a factorial design. Decomposition rates were similar between restoration treatments (p > 0.5), but leaves decomposed faster in the forest mixture than in the plantation mixture (p < 0.01). The content of Ca, Mg, K, P, and the C:N ratio were higher in the forest mixture at the beginning and during decomposition (p < 0.05); the N content in the plantation mixture was higher at the beginning but lower during decomposition (p < 0.05), which meant greater mobilization of nitrogen per unit of carbon lost. K and P had a strong initial release, while Mg was released more gradually. N and Ca had an irregular pattern of initial fast release, immobilization, and re‐release in the later stages. We conclude that the differences in rates of decomposition and nutrient release in these systems under restoration were at least partly determined by the floristic heterogeneity and chemical quality of the leaf litter that reaches the soil.     

The dynamics of surface acoustic wave‐driven scaffold cell seeding

Flow visualization using fluorescent microparticles and cell viability investigations are carried out to examine the mechanisms by which cells are seeded into scaffolds driven by surface acoustic waves. The former consists of observing both the external flow prior to the entry of the suspension into the scaffold and the internal flow within the scaffold pores. The latter involves micro‐CT (computed tomography) scans of the particle distributions within the seeded scaffolds and visual and quantitative methods to examine the morphology and proliferation ability of the irradiated cells. The results of these investigations elucidate the mechanisms by which particles are seeded, and hence provide valuable information that form the basis for optimizing this recently discovered method for rapid, efficient, and uniform scaffold cell seeding. Yeast cells are observed to maintain their size and morphology as well as their proliferation ability over 14 days after they are irradiated. The mammalian primary osteoblast cells tested also show little difference in their viability when exposed to the surface acoustic wave irradiation compared to a control set. Together, these provide initial feasibility results that demonstrate the surface acoustic wave technology as a viable seeding method without risk of denaturing the cells. Biotechnol. Bioeng. 2009;103: 387–401. © 2009 Wiley Periodicals, Inc.     

Climate‐driven reduction of genetic variation in plant phenology alters soil communities and nutrient pools

We examined the hypothesis that climate‐driven evolution of plant traits will influence associated soil microbiomes and ecosystem function across the landscape. Using a foundation tree species, Populus angustifolia, observational and common garden approaches, and a base population genetic collection that spans 17 river systems in the western United States, from AZ to MT, we show that (a) as mean annual temperature (MAT) increases, genetic and phenotypic variation for bud break phenology decline; (b) soil microbiomes, soil nitrogen (N), and soil carbon (C) vary in response to MAT and conditioning by trees; and (c) with losses of genetic variation due to warming, population‐level regulation of community and ecosystem functions strengthen. These results demonstrate a relationship between the potential evolutionary response of populations and subsequent shifts in ecosystem function along a large temperature gradient.     

Reconstructing the demise of Tethyan plants: climate‐driven range dynamics of Laurus since the Pliocene

Aim Climate changes are thought to be responsible for the retreat and eventual extinction of subtropical lauroid species that covered much of Europe and North Africa during the Palaeogene and early Neogene; little is known, however, of the spatial and temporal patterns of this demise. Herein we calibrate ecological niche models to assess the climatic requirements of Laurus L. (Lauraceae), an emblematic relic from the Tethyan subtropical flora, subsequently using these models to infer how the range dynamics of Laurus were affected by Plio‐Pleistocene climate changes. We also provide predictions of likely range changes resulting from future climatic scenarios. Location The Mediterranean Basin and Macaronesian islands (Canaries, Madeira, Azores). Methods We used a maximum‐entropy algorithm (Maxent) to model the relationship between climate and Laurus distribution over time. The models were fitted both to the present and to the middle Pliocene, based on fossil records. We employed climatic reconstructions for the mid‐Pliocene (3 Ma), the Last Glacial Maximum (21 ka) and a CO₂‐doubling future scenario to project putative species distribution in each period. We validated the model projections with Laurus fossil and present occurrences. Results Laurus preferentially occupied warm and moist areas with low seasonality, showing a marked stasis of its climatic niche. Models fitted to Pliocene conditions successfully predicted the current species distribution. Large suitable areas existed during the Pliocene, which were strongly reduced during the Pleistocene, but humid refugia within the Mediterranean Basin and Macaronesian islands enabled long‐term persistence. Future climate conditions are likely to re‐open areas suitable for colonization north of the current range. Main conclusions The climatic requirements of Laurus remained virtually unchanged over the last 3 Myr. This marked niche conservatism imposed largely deterministic range dynamics driven by climate conditions. This species's relatively high drought tolerance might account for the survival of Laurus in continental Europe throughout the Quaternary whilst other Lauraceae became extinct. Climatic scenarios for the end of this century would favour an expansion of the species's range towards northern latitudes, while severely limiting southern populations due to increased water stress.     

Integration of microbial kinetics and fluid dynamics toward model‐driven scale‐up of industrial bioprocesses

Scale‐up of bioprocesses is hampered by open questions, mostly related to poor mixing and mass transfer limitations. Concentration gradients of substrate, carbon dioxide, and oxygen in time and space, especially in large‐scale high‐cell density fed‐batch processes, are likely induced as the mixing time of the fermentor is usually longer than the relevant cellular reaction time. Cells in the fermentor are therefore repeatedly exposed to dynamic environments or perturbations. As a consequence, the heterogeneity in industrial practices often decreases either yield, titer, or productivity, or combinations thereof and increases by‐product formation as compared to well‐mixed small‐scale bioreactors, which is summarized as scale‐up effects. Identification of response mechanisms of the microorganism to various external perturbations is of great importance for pinpointing metabolic bottlenecks and targets for metabolic engineering. In this review, pulse response experimentation is proposed as an ideal way of obtaining kinetic information in combination with scale‐down approaches for in‐depth understanding of dynamic response mechanisms. As an emerging tool, computational fluid dynamics is able to draw a holistic picture of the fluid flow and concentration fields in the fermentor and finds its use in the optimization of fermentor design and process strategy. In the future, directed strain improvement and fermentor redesign are expected to largely depend on models, in which both microbial kinetics and fluid dynamics are thoroughly integrated.     

Rain,rats and pythons: Climate‐driven population dynamics of predators and prey in tropical Australia

Abstract: All natural populations fluctuate in abundance and age structure through time; understanding why they do so is a critical step towards their effective management and conservation. However, the long‐term data sets needed for such an understanding are rarely available, especially for tropical organisms. A 17‐year capture‐mark–recapture study yielded detailed information on the demography of water pythons (Liasis fuscus) and their main prey, the dusky rat (Rattus colletti), on the Adelaide River flood plain in tropical Australia. The link between annual rainfall patterns and rat demography was highly non‐linear. Rat numbers were low during years with low and high rainfall at the end of the wet season (April). Numbers of both predators and prey fluctuated considerably among years. Annual fluctuations in rat numbers generated a corresponding variation in rates of female python reproduction, python body condition and survival. Although variation in recruitment, survival and prey abundance all had a significant impact on annual fluctuations in python numbers, our analyses suggest that recruitment constituted the main determinant in driving the population dynamics of these large tropical predators. In combination with our other studies on this system, the data show that population dynamics of the water python population is ultimately driven by annual variation in rainfall, mediated via shifts in prey availability. The water pythons and the dusky rats of the Adelaide River flood plain thus demonstrate an unusually clear and direct link between an abiotic factor (rainfall) and predator–prey population dynamics.     

Diversity and abundance of human‐pathogenic fungi associated with pigeon faeces in urban environments

Pathogenic fungi are a growing health concern worldwide, particularly in large, densely populated cities. The dramatic upsurge of pigeon populations in cities has been implicated in the increased incidence of invasive fungal infections. In this study, we used a culture‐independent, high‐throughput sequencing approach to describe the diversity of clinically relevant fungi (CRF) associated with pigeon faeces and map the relative abundance of CRF across Seoul, Korea. In addition, we tested whether certain geographical, sociological and meteorological factors were significantly associated with the diversity and relative abundance of CRF. Finally, we compared the CRF diversity of fresh and old pigeon faeces to identify the source of the fungi and the role of pigeons in dispersal. Our results demonstrated that both the composition and relative abundance of CRF are unevenly distributed across Seoul. The green area ratio and the number of multiplex houses were positively correlated with species diversity, whereas wind speed and number of households were negatively correlated. The number of workers and green area ratio were positively correlated with the relative abundance of CRF, whereas wind speed was negatively correlated. Because many CRF were absent in fresh faeces, we inferred that most species cannot survive the gastrointestinal tract of pigeons and instead are likely transmitted through soil or air and use pigeon faeces as a substrate for proliferation.     

Spatial heterogeneity and short‐term oxygen dynamics in the rhizosphere of Vallisneria spiralis: Implications for nutrient cycling

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Plant invasion is associated with higher plant–soil nutrient concentrations in nutrient‐poor environments

Plant invasion is an emerging driver of global change worldwide. We aimed to disentangle its impacts on plant–soil nutrient concentrations. We conducted a meta‐analysis of 215 peer‐reviewed articles and 1233 observations. Invasive plant species had globally higher N and P concentrations in photosynthetic tissues but not in foliar litter, in comparison with their native competitors. Invasive plants were also associated with higher soil C and N stocks and N, P, and K availabilities. The differences in N and P concentrations in photosynthetic tissues and in soil total C and N, soil N, P, and K availabilities between invasive and native species decreased when the environment was richer in nutrient resources. The results thus suggested higher nutrient resorption efficiencies in invasive than in native species in nutrient‐poor environments. There were differences in soil total N concentrations but not in total P concentrations, indicating that the differences associated to invasive plants were related with biological processes, not with geochemical processes. The results suggest that invasiveness is not only a driver of changes in ecosystem species composition but that it is also associated with significant changes in plant–soil elemental composition and stoichiometry.     

Climate‐driven fluctuations in surface‐water availability and the buffering role of artificial pumping in an African savanna: Potential implication for herbivore dynamics

Abstract In arid and semiarid environments surface‐water strongly constrains the distribution and abundance of large herbivores during the dry season. Surprisingly, we know very little about its variability in natural ecosystems. Here we used long‐term data on the dry‐season occurrence of water at individual waterholes to model the surface‐water availability across Hwange National Park, Zimbabwe, under contrasted climatic and management scenarios. Without artificial pumping only 19.6% of the park occurred within 5 km of water under average climatic conditions. However surface‐water availability was strongly influenced by annual rainfall, and over 20 years the variability of the surface area of the park occurring within 5 km of water was slightly larger than the variability of rainfall. This contrasts with the usual buffered response of vegetation production to rainfall fluctuations, and suggests that the variability in dry‐season foraging range determined by surface‐water availability could be the main mechanism regulating the population dynamics of large herbivores in this environment. Artificial pumping increased surface‐water availability and reduced its variability over time. Because changes in surface‐water availability could cause the greatest changes in forage availability for large herbivores, we urge ecologists to investigate and report on the variability of surface‐water in natural ecosystems, particularly where rapid climate changes are expected.