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Lake Superior has recently begun experiencing cyanobacterial blooms comprised of Dolichospermum lemmermannii near the Apostle Islands and along the southern shore of the western arm. Little is known about the origin of these blooms. Experiments were conducted during the summers of 2017 and 2018 to identify sources of propagules and characteristics of sites that were potential sources. The 2017 experiments were conducted using a factorial design with three source zones ('River', 'Lake', and 'Harbor'), two nutrient conditions (high and low N:P), and three temperatures (15, 20, and 25°C). At the end of the experiment, cyanobacteria were most abundant from the 'River' and 'Harbor' zones at low N:P and 20 and 25°C, with D. lemmermannii most abundant at 20°C. Subsequently, in 2018 we evaluated 26 specific inland locations from three waterbody types ('River', 'Lake/Pond', and 'Coastal') and explored similarities among those sites that produced cyanobacteria in high abundance when samples were incubated under optimal conditions (low N:P and 25°C). Under these growing conditions, we found high cyanobacteria abundance developed in samples from river sites with low ambient temperatures and high conductivity. Field monitoring showed that Lake Superior nearshore temperatures were higher than rivers. These observations suggest that blooms of D. lemmermannii in Lake Superior are initiated by fluvial seeding of propagules and highlight the importance of warmer temperatures and favorable nutrient and light conditions for subsequent extensive cyanobacterial growth. We argue that the watershed is an important source of biological loading of D. lemmermannii to Lake Superior and that when those cells reach the nearshore where there are warmer water temperatures and increased light, they can grow in abundance to produce blooms., (Copyright © 2020. Published by Elsevier B.V.)

Winter conditions are rapidly changing in temperate ecosystems, particularly for those that experience periods of snow and ice cover. Relatively little is known of winter ecology in these systems, due to a historical research focus on summer 'growing seasons'. We executed the first global quantitative synthesis on under-ice lake ecology, including 36 abiotic and biotic variables from 42 research groups and 101 lakes, examining seasonal differences and connections as well as how seasonal differences vary with geophysical factors. Plankton were more abundant under ice than expected; mean winter values were 43.2% of summer values for chlorophyll a, 15.8% of summer phytoplankton biovolume and 25.3% of summer zooplankton density. Dissolved nitrogen concentrations were typically higher during winter, and these differences were exaggerated in smaller lakes. Lake size also influenced winter-summer patterns for dissolved organic carbon (DOC), with higher winter DOC in smaller lakes. At coarse levels of taxonomic aggregation, phytoplankton and zooplankton community composition showed few systematic differences between seasons, although literature suggests that seasonal differences are frequently lake-specific, species-specific, or occur at the level of functional group. Within the subset of lakes that had longer time series, winter influenced the subsequent summer for some nutrient variables and zooplankton biomass., (© 2016 The Authors. Ecology Letters published by CNRS and John Wiley & Sons Ltd.)

Human activities have increased the availability of reactive nitrogen in many ecosystems, leading to negative impacts on human health, biodiversity, and water quality. Freshwater ecosystems, including lakes, streams, and wetlands, are a large global sink for reactive nitrogen, but factors that determine the efficacy of freshwater nitrogen removal rates are poorly known. Using a global lake data set, we show that the availability of phosphorus, a limiting nutrient, affects both annual nitrogen removal rate and efficiency. This result indicates that increased phosphorus inputs from human activities have stimulated nitrogen removal processes in many lakes. Recent management-driven reductions in phosphorus availability promote water column accumulation and export of nitrogen from large lakes, an unintended consequence of single-element management that argues for greater control of nitrogen as well as phosphorus sources.

Disaccord between the supply and demand of energy (carbon, C) and certain material elements (e.g. phosphorus, P) across trophic levels is common in most ecosystems and impacts the strength of trophic interactions and ecosystem functions such as productivity and nutrient recycling. Yet, we know little about mechanisms operating at the lower levels of biological organization that drive such higher-level ecological processes. Such information should help refine theories integrating biological processes at multiple levels of organization. Understanding the expression and functions of genes that underlie (to a large degree) physiological adjustments made by organisms to stoichiometric imbalances at trophic interfaces is a first step in this enterprise. Here, we investigate adjustments in gene expression to varying supply and demand of phosphorus relative to other dietary components in the keystone limnetic herbivore, Daphnia pulex. Daphniids were fed an algal diet of either LoC-HiP (molar C:P ∼100) or HiC-LoP (molar C:P ∼900) for 5 days, resulting in significant growth reductions under HiC-LoP conditions. Microarrays measured the transcriptional regulation of 8217 annotated protein-coding genes under contrasting dietary conditions and revealed 1818 differentially expressed (DE) genes; 19% are genes unique to the Daphnia lineage. We mapped DE genes onto a global chart of metabolic pathways to obtain a systems-level perspective on the responses to stoichiometric imbalances. Daphnia differentially regulated pathways were involved in sequestering limiting elements, and in dealing with the products of metabolic adjustments that may be triggered by nutrient stress in primary producers. Functional genomics at trophic interfaces illuminate the complexity of processes underlying stoichiometric constraints on energy and nutrient fluxes in ecosystems., (© 2011 Blackwell Publishing Ltd.)

Multicellular organisms that benefit from division of labour are presumably descended from colonial species that initially derived benefits from larger colony size, before the evolution of specialization. Life in a colony can have costs as well as benefits, but these can be hard to measure. We measured physiological costs to life in a colony using a novel method based on population dynamics, comparing growth rates of unicells and kairomone-induced colonies of a green alga Desmodesmus subspicatus against a reference co-occurring species. Coloniality negatively affected growth during the initial log growth phase, while no adverse effect was detected under nutrient-limited competitive conditions. The results point to costs associated with traits involved in rapid growth rather than those associated with efficient growth under resource scarcity. Some benefits of coloniality (e.g. defence from herbivory) may be different from when this trait evolved, but our approach shows how costs would have depended on conditions.

The genetic basis of organism response to stoichiometric mismatches between environmental availability and somatic demand is still poorly understood. This study reports a consistent genotype x environment interaction related to phosphorus : carbon availability to Daphnia. In multiple pairs of Daphnia pulicaria clones, genetic variation at the phosphoglucose isomerase (Pgi) locus indicated that Pgi-heterozygotes out competed Pgi-homozygotes under high P : C conditions, whereas the opposite outcome was observed under low P : C conditions. Estimates of phosphorus use efficiency indicated that homozygotes were significantly more efficient. However, homozygotes were comparatively less homeostatic. We hypothesize that lower specific activity of Pgi from homozygotes, which results in lowered energetic efficiency during the second glycolytic step, may underlie the competitive advantage enjoyed by homozygotes under low P : C (i.e. excess C) conditions. Our results show that analysing stoichiometric mismatches between diet and consumer should advance our quest for a fundamental understanding of the mechanisms driving genotype-environment interactions.

A century-long increase in nitrate (NO3-) in the water column of Lake Superior is a classic example of recent nitrogen accumulation in ecosystems, but its cause and relationship to historical NO3- deposition is unknown. We used stable isotope ratios of oxygen and nitrogen in nitrate (delta18O-NO3 and delta15N-NO3) to examine its sources in this large lake, which represents 10% of the world's surficial liquid freshwater. The most parsimonious hypothesis to explain the rise in NO3- is that the lake is accruing NO3- deposited directly on the lake surface because it is too unproductive to completely assimilate all of it. Data for delta18O-NO3 in external sources and the water column, however, are inconsistent with this hypothesis. Instead, the isotopic evidence indicates strongly that the accumulating NO3- is almost entirely derived from nitrification occurring within the lake. While increases in atmospheric deposition of NO3- may have played a role in its buildup in the lake, other factors such as increases in NH4+ and dissolved organic nitrogen inputs from precipitation or rivers, increases in nitrogen fluxes from the sediments, and decreases in burial rates must also be considered as potential drivers of rising NO3-. The sustained accumulation of nitrogen in Lake Superior is thus more complex and incompletely understood than previously assumed.

Biological and environmental contrasts between aquatic and terrestrial systems have hindered analyses of community and ecosystem structure across Earth's diverse habitats. Ecological stoichiometry provides an integrative approach for such analyses, as all organisms are composed of the same major elements (C, N, P) whose balance affects production, nutrient cycling, and food-web dynamics. Here we show both similarities and differences in the C:N:P ratios of primary producers (autotrophs) and invertebrate primary consumers (herbivores) across habitats. Terrestrial food webs are built on an extremely nutrient-poor autotroph base with C:P and C:N ratios higher than in lake particulate matter, although the N:P ratios are nearly identical. Terrestrial herbivores (insects) and their freshwater counterparts (zooplankton) are nutrient-rich and indistinguishable in C:N:P stoichiometry. In both lakes and terrestrial systems, herbivores should have low growth efficiencies (10-30%) when consuming autotrophs with typical carbon-to-nutrient ratios. These stoichiometric constraints on herbivore growth appear to be qualitatively similar and widespread in both environments.

The amounts of solar energy and materials are two of the chief factors determining ecosystem structure and process. Here, we examine the relative balance of light and phosphorus in a set of freshwater pelagic ecosystems. We calculated a ratio of light: phosphorus by putting mixed-layer mean light in the numerator and total P concentration in the denominator. This light: phosphorus ratio was a good predictor of the C:P ratio of particulate matter (seston), with a positive correlation demonstrated between these two ratios. We argue that the balance between light and nutrients controls "nutrient use efficiency" at the base of the food web in lakes. Thus, when light energy is high relative to nutrient availability, the base of the food web is carbon rich and phosphorus poor. In the opposite case, where light is relatively less available compared to nutrients, the base of the food web is relatively P rich. The significance of this relationship lies in the fact that the composition of sestonic material is known to influence a large number of ecosystem processes such as secondary production, nutrient cycling, and (we hypothesize) the relative strength of microbial versus grazing processes. Using the central result of increased C:P ratio with an increased light: phosphorus ratio, we make specific predictions of how ecosystem structure and process should vary with light and nutrient balance. Among these predictions, we suggest that lake ecosystems with low light: phosphorus ratios should have several trophic levels simultaneously carbon or energy limited, while ecosystems with high light: phosphorus ratios should have several trophic levels simultaneously limited by phosphorus. Our results provide an alternative perspective to the question of what determines nutrient use efficiency in ecosystems.

Experiments using planktonic organisms revealed that the balance of radiant energy and available nutrients regulated herbivore growth rates through their effects on abundance and chemical composition of primary producers. Both algae and herbivores were energy limited at low light/nutrient ratios, but both were nutrient limited at high light/nutrient ratios. Herbivore growth increased with increasing light intensity at low values of the light/nutrient ratio due to increases in algal biomass, but growth decreased with increasing light at a high light/nutrient ratio due to decreases in algal quality. Herbivore production therefore was maximal at intermediate levels of the light/nutrient ratio. The results contribute to an understanding of mass transfer mechanisms in ecosystems and illustrate the importance of integration of energy-based and material-based currencies in ecology.

Bacterial abundance results from predatory losses of individuals and replacement of losses through growth. Growth depends on sustained input of organic substrates and mineral nutrients. In this work we tested the hypothesis that bacterial growth in two oligotrophic Canadian shield lakes was limited by nitrogen (N) or phosphorus (P). We also determined whether consumer-regenerated resources contributed substantially to net bacterial growth. Two types of dilution assays were conducted to determine the response of bacteria to nutrient enrichment: diluted whole water (DWW, 1:9 whole/filtered with 0.2 μm of filtered lake water) and diluted fractionated water (DFW, 1.0 μm prefiltered then diluted as above). Replicate bottles in each dilution assay received either N (50 μM), P (10 μM), or both N and P enrichments. Controls received no nutrients. Resource-saturated growth rates and grazing rates were estimated from a standard dilution-growth approach. Bacterial growth was stimulated by addition of P alone and in combination with N. Consumers regenerated sufficient resources to support up to half the bacterial growth rate, but the benefit derived from consumers was minor when compared to mortality.

The dynamics of carbon (C), nitrogen (N), and phosphorus (P), elemental ratios, and dark uptake/release of N and P in bacterial and phytoplankton size fractions were studied during summer 1992 in three lakes of contrasting food web structure and trophic status (L240, L110, L227). We wished to determine if phytoplankton and bacteria differed in their elemental characteristics and to evaluate whether the functional role of bacteria in nutrient cycling (i.e., as sink or source) depended on bacterial elemental characteristics. Bacterial contributions to total suspended particulate material and to fluxes of nutrients in the dark were substantial and varied for different elements. This indicated that some techniques for assaying phytoplankton physiological condition are compromised by bacterial contributions. C/N ratios were generally less variable than C/P and N/P ratios. Both elemental ratios and biomass-normalized N and P flux indicated that phytoplankton growth in each lake was predominantly P-limited, although in L227 these data reflect the dominance of N-fixing cyanobacteria, and N was likely limiting early in the sampling season. In L227, phytoplankton N/P ratio and biomass-normalized N flux were negatively correlated, indicating that flux data were likely a reasonable measure of the N status of the phytoplankton. However, for L227 phytoplankton, P-flux per unit biomass was a hyperbolic function of N/P, suggesting that the dominant L227 cyanobacteria have a limited uptake and storage capacity and that P-flux per unit biomass may not be a good gauge of the P-limitation status of phytoplankton in this situation. Examination of N-flux data in the bacterial size fraction relative to the N/P ratio of the bacteria revealed a threshold N/P ratio (∼22:1 N/P, by atoms), below which, bacteria took up and sequestered added N, and above which, N was released. Thus, the functional role of bacteria in N cycling in these ecosystems depended on their N/P stoichiometry.

Single-species, steady-state chemostat cultures of two freshwater diatoms showed that Fragilaria crotonensis had a lower equilibrial requirement for silicate than did Tabellaria fenestratra. Using this information, resource competition theory predicts that Fragilaria should be able to invade silicate-limited equilibrium populations of Tabellaria, but Tabellaria should not be able to invade silicatelimited Fragilaria. This prediction was supported by a series of invasion experiments. The two species did not have detectable differences in their phosphate requirements. Invasion experiments showed that Fragilaria could invade Tabellaria cultures, but that Tabellaria could not invade Fragilaria cultures under phosphate-limited conditions. These results are consistent with predictions based on previous studies of the phosphate physiology of these genera.

The increase in algal reproductive rates caused by nitrogen regeneration from herbivorous zooplankton approximately equaled the zooplankton-caused mortality. This result demonstrates that nutrient regeneration by herbivores is at least sometimes a strong indirect effect in natural communities.