Dynamic responses of phosphorus metabolism to acute and chronic dietary phosphorus-limitation in Daphnia

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 rates 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 h. 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

Seasonal effects of food quality and temperature on body stoichiometry, biochemistry, and biomass production in Daphnia populations

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

Interactive effects of genotype and food quality on consumer growth rate and elemental content

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

Fear and food: Effects of predator-derived chemical cues and stoichiometric food quality on Daphnia

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

The threshold elemental ratio of carbon and phosphorus of Daphnia magna and its connection to animal growth

The growth of animal consumers is afected by the balance of elements in their diet with the transition between limitation by one element to another known as the threshold elemental ratio (TER). Precise estimates of TERs with known levels of uncertainty have yet to be generated for most zooplankton consumers. We determined the TER for carbon (C) and phosphorus (P) in for a common lake zooplankter, Daphnia magna, using experimental measurements and theoretical considerations. Daphnia growth responses to food C:P ratios across a relatively narrow range (80–350) generated an empirical estimate of TERC:P of 155±14. While this TER matched our modelled estimate of TERC:P (155±16), it was lower than previous estimates of this dietary transition point. No threshold was found when we examined daphnid body C:N or C:P ratios in response to changing food C:P ratios, which indicates P-limitation at even lower food C:P ratios. Our results provide strong evidence that D. magna is likely to experience acute P-limitation when food C:P ratios exceed even relatively low ratios (~155). Our model further demonstrated that while physiological adjustments may reduce the likelihood of P-limitation or reduce its intensity, these changes in animal material processing would be accompanied by reduced maximum growth rates

Mobility and bioavailability of sediment phosphorus in urban stormwater ponds

© 2019. American Geophysical Union. All Rights Reserved. Stormwater ponds can serve as retention hotspots for phosphorus (P) moving out of the urban environment. This retention may be reduced by P speciation that reduces the bioavailability of P to primary producers and alters its mobility in sediments. Here we examined the mobility and fate of dissolved P in urban stormwater ponds with a set of complementary field measurements and short-term laboratory and field experiments. We measured the types and amount of P in water column and sediments of urban stormwater ponds. We further assessed the mobility of different P types in pond sediments in the field and rates of P release from sediment cores maintained under laboratory conditions. Finally, we assessed P uptake rates by pond algal communities using short-term bioassay experiments. We found that dissolved organic P was highly prevalent in urban pond water and sediments and that this type of P was mobile within sediments and could be released under high or low O 2 conditions. We also found highly variable P demand by algae among stormwater ponds and that algal growth responses to P was correlated to water column N:P ratios. Altogether, our results indicate an important role for organic phosphorus cycling in urban stormwater ponds, which likely constrains the overall retention efficiency in these aquatic ecosystems

Variation in particulate C : N : P stoichiometry across the Lake Erie watershed from tributaries to its outflow

© 2017 The Authors Limnology and Oceanography published by Wiley Periodicals, Inc. on behalf of Association for the Sciences of Limnology and Oceanography 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

Effects of calcium and phosphorus limitation on the nutritional ecophysiology of Daphnia

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

Appendix A. Seven component PARAFAC model loadings and split-half validation.

Seven component PARAFAC model loadings and split-half validation

Phosphorus Availability Alters the Effects of Silver Nanoparticles on Periphyton Growth and Stoichiometry

<div><p>Exposure to silver nanoparticles (AgNPs) may alter the structure and function of freshwater ecosystems. However, there remains a paucity of studies investigating the effects of AgNP exposure on freshwater communities in the natural environment where interactions with the ambient environment may modify AgNP toxicity. We used nutrient diffusing substrates to determine the interactive effects of AgNP exposure and phosphorus (P) enrichment on natural assemblages of periphyton in three Canadian Shield lakes. The lakes were all phosphorus poor and spanned a gradient of dissolved organic carbon availability. Ag slowly accumulated in the exposed periphyton, which decreased periphyton carbon and chlorophyll <i>a</i> content and increased periphyton C:P and N:P in the carbon rich lakes. We found significant interactions between AgNP and P treatments on periphyton carbon, autotroph standing crop and periphyton stoichiometry in the carbon poor lake such that P enhanced the negative effects of AgNPs on chlorophyll <i>a</i> and lessened the impact of AgNP exposure on periphyton stoichiometry. Our results contrast with those of other studies demonstrating that P addition decreases metal toxicity for phytoplankton, suggesting that benthic and pelagic primary producers may react differently to AgNP exposure and highlighting the importance of <i>in situ</i> assays when assessing potential effects of AgNPs in fresh waters.</p></div