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Human activities can cause large alterations in biogeochemical cycles of key nutrients such as carbon (C), nitrogen (N), and phosphorus (P). However, relatively little is known about how these changes alter the proportional fluxes of these elements across ecosystem boundaries from rivers to lakes. Here, we examined environmental factors influencing spatial and temporal variation in particulate C:N:P ratios across the Lake Erie watershed from its tributaries to its outflow. Throughout the study, particulate nutrient ratios ranged widely (C:N 2.0–25.8, C:P 32–530, N:P 3.7–122.9), but mean values were generally lower than previous estimates from different aquatic environments. Particulate C:N ratios varied the least across all environments, but C:P and N:P ratios increased between tributaries and coastal areas and throughout the growing season in coastal environments. These ratios also differed temporally in offshore waters as particulate C:P and N:P were higher in the spring and summer and lower in the fall and winter. Particulate C:P ratios also increased between the western/central and eastern basins indicating differential nutrient processing across the lake. These stoichiometric changes were associated with unique environmental factors among ecosystems as tributary stoichiometry was related to terrestrial land use and land cover, coastal ratios were a product of mixing between riverine and offshore waters, and offshore patterns were influenced by differences in temperature and particulate nutrient loading among basins. Overall, by studying changes in particulate C:N:P ratios across the Lake Erie watershed, our study demonstrates the power of using mass balance principles to study nutrient transformations along the aquatic continuum.

While the sentinel nature of freshwater systems is now well recognized, widespread integration of freshwater processes and patterns into our understanding of broader climate-driven Arctic terrestrial ecosystem change has been slow. We review the current understanding across Arctic freshwater systems of key sentinel responses to climate, which are attributes of these systems with demonstrated and sensitive responses to climate forcing. These include ice regimes, temperature and thermal structure, river baseflow, lake area and water level, permafrost-derived dissolved ions and nutrients, carbon mobilization (dissolved organic carbon, greenhouse gases, and radiocarbon), dissolved oxygen concentrations, lake trophic state, various aquatic organisms and their traits, and invasive species. For each sentinel, our objectives are to clarify linkages to climate, describe key insights already gained, and provide suggestions for future research based on current knowledge gaps. We suggest that tracking key responses in Arctic freshwater systems will expand understanding of the breadth and depth of climate-driven Arctic ecosystem changes, provide early indicators of looming, broader changes across the landscape, and improve protection of freshwater biodiversity and resources.

Recent studies have shown that adaptive evolution can be rapid enough to affect contemporary ecological dynamics in nature (i.e. ‘rapid evolution’). These studies tend to focus on trait functions relating to interspecific interactions; however, the importance of rapid evolution of stoichiometric traits has been relatively overlooked. Various traits can affect the balance of elements (carbon, nitrogen, and phosphorus) of organisms, and rapid evolution of such stoichiometric traits will not only alter population and community dynamics but also influence ecosystem functions such as nutrient cycling. Multiple environmental changes may exert a selection pressure leading to adaptation of stoichiometrically important traits, such as an organism's growth rate. In this paper, we use theoretical approaches to explore the connections between rapid evolution and ecological stoichiometry at both the population and ecosystem level. First, we incorporate rapid evolution into an ecological stoichiometry model to investigate the effects of rapid evolution of a consumer's stoichiometric phosphorus:carbon ratio on consumer–producer population dynamics. We took two complementary approaches, an asexual clonal genotype model and a quantitative genetic model. Next, we extended these models to explicitly track nutrients in order to evaluate the effect of rapid evolution at the ecosystem level. Our model results indicate rapid evolution of the consumer stoichiometric trait can cause complex dynamics where rapid evolution destabilizes population dynamics and rescues the consumer population from extinction (evolutionary rescue). The model results also show that rapid evolution may influence the level of nutrients available in the environment and the flux of nutrients across trophic levels. Our study represents an important step for theoretical integration of rapid evolution and ecological stoichiometry.

Food quality is highly dynamic within lake ecosystems and varies spatially and temporally over the growing season. Consumers may need to continuously adjust their metabolism in response to this variation in dietary nutrient content. However, the rate of metabolic responses to changes in food nutrient content has received little direct study. Here, we examine responses in two metabolic phosphorus (P) pools, ribonucleic acids (RNA), and adenosine triphosphate (ATP) along with body mass and body P content in Daphnia magna exposed to chronic and acute dietary P-limitation. First, we examined food quality effects on animals consuming different food carbon (C):P quality over a 14 day period. Then, we raised daphnids on one food quality for 4 days, switched them to contrasting dietary treatments, and measured changes in their metabolic responses at shorter time-scales (over 48 h). Animal P, RNA, and ATP content all changed through ontogeny with adults containing relatively less of these pools with increasing body mass. Irrespective of age, Daphnia consuming high C:P diets had lower body %P, %RNA, %ATP, and mass compared to animals eating low C:P diets. Diet switching experiments revealed diet dependent changes in body %P, %RNA, %ATP, and animal mass within 48 hours. We found that Daphnia switched from low to high C:P diets had some metabolic buffering capacity with decreases in body %P occurring after 24 h but mass remaining similar to initial diet conditions for 36 h after the diet switch. Switching Daphnia from low to high C:P diets caused a decrease in the RNA:P ratio after 48 h. Daphnia switched from high to low C:P diets increased their body P, RNA, and ATP content within 8-24 h. This switch from high to low C:P diets also led to increased RNA:P ratios in animal bodies. Overall, our study revealed that consumer P metabolism reflects both current and past diet due to more dynamic and rapid changes in P biochemistry than total body mass. This metabolic flexibility is likely linked to resource integration in D. magna, which reduces the negative effects of short-term or variable exposure to nutrient-deficient foods.

Food quality and temperature can affect zooplankton production in lakes by altering organismal metabolism. However, the influence of these factors on consumer nutritional physiology and population biomass remains relatively understudied in natural populations. Here, we examined seasonal changes in body stoichiometry, biochemistry, and population biomass in two Daphnia species collected from two separate lakes differing in dietary phosphorus (P) supply. Food quality, measured as seston carbon:P (C:P) ratios, varied throughout the study in each lake, and water temperatures generally increased across the growing season. Daphnid elemental composition was correlated with food quality in both populations, but relationships between daphnid body stoichiometry and temperature were consistently stronger as Daphnia body C:P ratios and content of major biochemical pools declined simultaneously throughout the summer, which largely coincided with increased water temperatures. Warmer temperatures were associated with relaxed %P-RNA coupling as daphnid body RNA content declined and P content remained relatively high. These responses combined with temperature related decreases in Daphnia body %lipids and %C appeared to explain declines in daphnid body C:P ratios in both lakes over the growing season. Seasonal changes in population biomass were related to both food quality and water temperature in the lower nutrient lake. Biomass production under more eutrophic conditions however was unrelated to food quality and was instead associated with seasonal temperature changes in the higher nutrient lake. Overall, our study shows that seasonal changes in temperature and resource quality may differentially affect consumer stoichiometry and biomass production in lake ecosystems by altering consumer elemental metabolism.

While resource quality and predator-derived chemical cues can each have profound effects on zooplankton populations and their function in ecosystems, the strength and direction of their interactive effects remain unclear. We conducted laboratory experiments to evaluate how stoichiometric food quality (i.e., algal carbon [C] : phosphorus [P] ratios) affects responses of the zooplankter, Daphnia pulicaria, to predator-derived chemical cues. We compared growth rates, body P content, metabolic rates, life-history shifts, and survival of differentially P-nourished Daphnia in the presence and absence of chemical cues derived from fish predators. We found effects of predator cues and/or stoichiometric food quality on all measured traits of Daphnia. Exposure to fish cues led to reduced growth and increased metabolic rates but had little effect on the body %P content of Daphnia. Elevated algal C : P ratios reduced growth and body %P and increased mass-specific respiration rates. While most of the effects of predator cues and algal C : P ratios of Daphnia were non-interactive, reduced survival and relatedly reduced population growth rates that resulted from P-poor food were amplified in the presence of predator-derived cues. Our results demonstrate that stoichiometric food quality interacts with antipredator responses of Daphnia, but these effects are largely trait dependent and appear connected to animal life-history evolution. Given the ubiquity of predators and P-poor food in lake ecosystems, our results highlight the importance of the interactive responses of animals to predator cues and poor nutrition.

Consumer body stoichiometry is a key trait that links organismal physiology to population and ecosystem-level dynamics. However, as elemental composition has traditionally been considered to be constrained within a species, the ecological and evolutionary factors shaping consumer elemental composition have not been clearly resolved. To this end, we examined the causes and extent of variation in the body phosphorus (P) content and the expression of P-linked traits, mass specific growth rate (MSGR), and P use efficiency (PUE) of the keystone aquatic consumer Daphnia using lake surveys and common garden experiments. While daphnid body %P was relatively constrained in field assemblages sampled across an environmental P gradient, unique genotypes isolated from these lakes showed highly variable phenotypic responses when raised across dietary P gradients in the laboratory. Specifically, we observed substantial inter- and intra-specific variation and differences in daphnid responses within and among our study lakes. While variation in Daphnia body %P was mostly due to plastic phenotypic changes, we documented considerable genetic differences in daphnid MSGR and PUE, and relationships between MSGR and body P content were highly variable among genotypes. Overall, our study found that consumer responses to food quality may differ considerably among genotypes and that relationships between organismal life-history traits and body stoichiometry may be strongly influenced by genetic and environmental variation in natural assemblages.

Declines in environmental calcium (Ca) and phosphorus (P) concentrations have occurred over the past 30 yrs in lakes across the Canadian Shield in southern Ontario, and these reductions appear to be placing strong constraints on populations of Daphnia in this region. Here, we report results from a factorial manipulation of Ca concentrations and food P content under controlled laboratory conditions where we measured resulting changes in daphnid elemental content, individual growth and survival, and life history traits related to population growth. We found significant effects of Ca- and P-limitation on all variables measured; however, dietary P explained a majority of the variation in daphnid nutrient content and growth. Dietary effects of low P [high food carbon (C): P ratios] on individual Daphnia life-history traits also translated into significant population level effects. Dietary P also explained relatively more experimental variation in population level responses than Ca concentrations. Experimental Ca concentrations most strongly altered daphnid survival partly due to the use of a lethally low Ca concentration in our experiment. Although recent work examining shifts in zooplankton communities in this region mainly focuses on the effects of Ca-limitation, we show that Ca concentrations and food nutrient content, at levels commonly found on the Canadian Shield, are both likely to strongly alter Daphnia life-history and populations dynamics. Our results underscore the need to more fully examine how multielemental limitation (e.g., Ca, N, P) affects consumer physiology and life-history given the plausible translation of these effects on the community structure of lake zooplankton.

Global change is reshaping the physical environment and altering nutrient dynamics across the Arctic. These changes can affect the structure and function of biological communities and influence important climate-related feedbacks (for example, carbon (C) sequestration) in biogeochemical processing hot spots such as lakes. To understand how these ecosystems will respond in the future, this study examined recent (< 10 y) and long-term (1000 y) shifts in autotrophic production across paraglacial environmental gradients in SW Greenland. Contemporary lake temperatures and light levels increased with distance from the ice sheet, along with dissolved organic C (DOC) concentrations and total nitrogen: total phosphorus (TN:TP) ratios. Diatom production measured as biogenic silica accumulation rates (BSiARs) and diatom contribution to microbial communities declined across these gradients, while total production estimated using C accumulation rates and δ¹³C increased, indicating that autochthonous production and C burial are controlled by microbial competition and competitive displacement across physiochemical gradients in the region. Diatom production was generally low across lakes prior to the 1800’s AD but has risen 1.5–3× above background levels starting between 1750 and 1880 AD. These increases predate contemporary regional warming by 115–250 years, and temperature stimulation of primary production was inconsistent with paleorecords for ~ 90% of the last millennium. Instead, primary production appeared to be more strongly related to N and P availability, which differs considerably across the region due to lake landscape position, glacial activity and degree of atmospheric nutrient deposition. These results suggest that biological responses to enhanced nutrient supply could serve as important negative feedbacks to global change.

Many lakes across Canada and northern Europe have experienced declines in ambient phosphorus (P) and calcium (Ca) supply for over 20 years. While these declines might create or exacerbate nutrient limitation in aquatic food webs, our ability to detect and quantify different types of nutrient stress on zooplankton remains rudimentary. Here, we used growth bioassay experiments and whole transcriptome RNAseq, collectively nutrigenomics, to examine the nutritional phenotypes produced by low supplies of P and Ca separately and together in the freshwater zooplankter Daphnia pulex. We found that daphniids in all three nutrient-deficient categories grew slower and differed in their elemental composition. Our RNAseq results show distinct responses in singly limited treatments (Ca or P) and largely a mix of these responses in animals under low Ca and P conditions. Deeper investigation of effect magnitude and gene functional annotations reveals this patchwork of responses to cumulatively represent a co-limited nutritional phenotype. Linear discriminant analysis identified a significant separation between nutritional treatments based upon gene expression patterns with the expression patterns of just five genes needed to predict animal nutritional status with 99% accuracy. These data reveal how nutritional phenotypes are altered by individual and co-limitation of two highly important nutritional elements (Ca and P) and provide evidence that aquatic consumers can respond to limitation by more than one nutrient at a time by differentially altering their metabolism. This use of nutrigenomics demonstrates its potential to address many of the inherent complexities in studying interactions between multiple nutritional stressors in ecology and beyond.