Phytoplankton Nutrient Limitation Research Articles

Global analysis of ocean phytoplankton nutrient limitation reveals high prevalence of co-limitation

Nutrient availability limits phytoplankton growth throughout much of the global ocean. Here we synthesize available experimental data to identify three dominant nutrient limitation regimes: nitrogen is limiting in the stratified subtropical gyres and in the summertime Arctic Ocean, iron is most commonly limiting in upwelling regions, and both nutrients are frequently co-limiting in regions in between the nitrogen and iron limited systems. Manganese can be co-limiting with iron in parts of the Southern Ocean, whilst phosphate and cobalt can be co-/serially limiting in some settings. Overall, an analysis of experimental responses showed that phytoplankton net growth can be significantly enhanced through increasing the number of different nutrients supplied, regardless of latitude, temperature, or trophic status, implying surface seawaters are often approaching nutrient co-limitation. Assessments of nutrient deficiency based on seawater nutrient concentrations and nutrient stress diagnosed via molecular biomarkers showed good agreement with experimentally-assessed nutrient limitation, validating conceptual and theoretical links between nutrient stoichiometry and microbial ecophysiology.

Persistent equatorial Pacific iron limitation under ENSO forcing.

Projected responses of ocean net primary productivity to climate change are highly uncertain1. Models suggest that the climate sensitivity of phytoplankton nutrient limitation in the low-latitude Pacific Ocean plays a crucial role1-3, but this is poorly constrained by observations4. Here we show that changes in physical forcing drove coherent fluctuations in the strength of equatorial Pacific iron limitation through multiple El Niño/Southern Oscillation (ENSO) cycles, but that this was overestimated twofold by a state-of-the-art climate model. Our assessment was enabled by first using a combination of field nutrient-addition experiments, proteomics and above-water hyperspectral radiometry to show that phytoplankton physiological responses to iron limitation led to approximately threefold changes in chlorophyll-normalized phytoplankton fluorescence. We then exploited the >18-year satellite fluorescence record to quantify climate-induced nutrient limitation variability. Such synoptic constraints provide a powerful approach for benchmarking the realism of model projections of net primary productivity to climate changes.

Co-Limitation of Phytoplankton by N and P in a Shallow Coastal Lagoon (Ria Formosa): Implications for Eutrophication Evaluation

AbstractThe Ria Formosa coastal lagoon is a highly productive shallow ecosystem in southern Portugal, subjected to nutrient inputs from anthropogenic and natural sources. Nutrients are major abiotic drivers of phytoplankton in this system, but their effects on phytoplankton assemblages and the occurrence of nutrient limitation are still poorly understood. The main goal of this study was, thus, to evaluate the occurrence, type, and effects of nutrient limitation on phytoplankton community and specific functional groups in the Ria Formosa coastal lagoon. We conducted nutrient enrichment experiments with factorial additions of nitrogen (N) and phosphorus (P) using natural phytoplankton assemblages from distinct locations in the Ria Formosa, throughout a yearly cycle. Phytoplankton composition and abundance were evaluated using inverted and epifluorescence microscopies, and spectrophotometric methods were used for biomass. Limitation was defined as higher phytoplankton growth following enrichment with a particular nutrient in relation to the non-enriched control. The most common type of phytoplankton limitation was simultaneous co-limitation by N and P; diatoms, as r-strategists, were the most frequently limited group. Single N and P limitation, and serial P limitation were also observed, as well as negative responses to nutrient enrichment. Group-specific responses to nutrient enrichment were not reflected in the relative abundance of phytoplankton groups within the whole assemblage, due to the numerical dominance of pico-sized groups (cyanobacteria and eukaryotic picophytoplankton). Ambient nutrient ratios and concentrations did not predict phytoplankton nutrient limitation, given the different nutrient utilisation traits among phytoplankton functional groups. Therefore, nutrient ratios should not be used as indicators of nutrient limitation in eutrophication assessment.

Regime shifts in shallow lakes explained by critical turbidity

Worldwide, water quality managers target a clear, macrophyte-dominated state over a turbid, phytoplankton-dominated state in shallow lakes. The competition mechanisms underlying these ecological states were explored in the 1990s, but the concept of critical turbidity seems neglected in contemporary water quality models. In particular, a simple mechanistic model of alternative stable states in shallow lakes accounting for resource competition mechanisms and critical turbidity is lacking. To this end, we combined Scheffer's theory on critical turbidity with insights from nutrient and light competition theory founded by Tilman, Huisman and Weissing. This resulted in a novel graphical and mathematical model, GPLake-M, that is relatively simple and mechanistically understandable and yet captures the essential mechanisms leading to alternative stable states in shallow lakes. The process-based PCLake model was used to parameterize the model parameters and to test GPLake-M using a pattern-oriented strategy. GPLake-M's application range and position in the model spectrum are discussed. We believe that our results support the fundamental understanding of regime shifts in shallow lakes and provide a starting point for further mechanistic and management-focused explorations and model development. Furthermore, the concept of critical turbidity and the relation between light-limited submerged macrophytes and nutrient-limited phytoplankton might provide a new focus for empirical aquatic ecological research and water quality monitoring programs.

Nitrogen is unlikely to consistently limit primary productivity in most tropical lakes

AbstractDespite mounting evidence that tropical lakes may experience an array of phytoplankton nutrient limitation regimes, references to low‐latitude lakes being predominantly nitrogen limited remain common in the literature. To assess the current understanding of nutrient limitation regimes in tropical lakes, we performed a literature review with the following objectives: First, quantify and summarize observations of nitrogen (N), phosphorus (P), and colimitation in the collected studies. Second, identify common drivers of tropical nutrient limitation regimes. Third, provide literature‐informed recommendations for future research efforts into nutrient limitation of tropical lakes. We report that not only is N not the dominant limiting nutrient in the reviewed lakes, but tropical lakes exhibit a wide range of nutrient limitation regimes, and there is often considerable heterogeneity of the dominant limiting nutrient within an individual lake. This heterogeneity is driven largely by seasonal patterns of water column stratification and precipitation, land use and land cover (LULC), and the interaction of these system characteristics with lake morphology. To accurately characterize nutrient limitation in tropical aquatic ecosystems, assessments need to include sampling across space and time in order to capture within‐lake spatial heterogeneity and both inter‐ and intra‐annual temporal variation. Whenever possible, both inferential and experimental approaches should be used in concert to determine nutrient limitation due to methodological limitations.

Nutrient function over form: Organic and inorganic nitrogen additions have similar effects on lake phytoplankton nutrient limitation

AbstractThe concentration of dissolved organic nitrogen (DON) is increasing in many northern hemisphere lakes, yet its use by phytoplankton and fate in the environment seldom have been quantified. We conducted 1 week, in situ, microcosm incubations across 25 lakes in northeastern North America to understand how DON, dissolved inorganic nitrogen (DIN), and dissolved inorganic phosphorus (P) affected phytoplankton biomass. In addition, we tested whether lakes were limited by single macronutrients (N or P) or colimited by both. Phytoplankton biomass in 80% of lakes responded similarly to DON and DIN additions. Of the lakes where N form produced differential responses, the majority of phytoplankton communities exhibited greater biomass accumulation with DON than DIN. Colimitation was the most common type of nutrient limitation among the study lakes, followed by P limitation. Limitation type shifted with N form in 40% of the study lakes, but without consistent patterns explaining how shifts occurred. Regardless of N form, lakes with watersheds more dominated by agriculture and higher total dissolved nitrogen (TDN) tended to show P‐limited phytoplankton responses, while lakes with less agricultural watersheds and lower TDN tended to show colimited phytoplankton responses. Finally, ambient TDN and total phosphorus (TP) nutrient concentrations were stronger predictors of limitation type than ambient TDN : TP ratios. The different contributions of DON and DIN to phytoplankton biomass in some of our study lakes suggest that DON loading from surrounding watersheds may be an overlooked component in predicting phytoplankton productivity and nutrient limitation dynamics in freshwater ecosystems.

Active reconfiguration of cytoplasmic lipid droplets governs migration of nutrient-limited phytoplankton.

Nutrient availability, along with light and temperature, drives marine primary production. The ability to migrate vertically, a critical trait of motile phytoplankton, allows species to optimize nutrient uptake, storage, and growth. However, this traditional view discounts the possibility that migration in nutrient-limited waters may be actively modulated by the emergence of energy-storing organelles. Here, we report that bloom-forming raphidophytes harness energy-storing cytoplasmic lipid droplets (LDs) to biomechanically regulate vertical migration in nutrient-limited settings. LDs grow and translocate directionally within the cytoplasm, steering strain-specific shifts in the speed, trajectory, and stability of swimming cells. Nutrient reincorporation restores their swimming traits, mediated by an active reconfiguration of LD size and coordinates. A mathematical model of cell mechanics establishes the mechanistic coupling between intracellular changes and emergent migratory behavior. Amenable to the associated photophysiology, LD-governed behavioral shift highlights an exquisite microbial strategy toward niche expansion and resource optimization in nutrient-limited oceans.

Phytoplankton community structuring in the absence of resource-based competitive exclusion.

Under most natural marine conditions, phytoplankton cells suspended in the water column are too distantly spaced for direct competition for resources (i.e., overlapping cell boundary layers) to be a routine occurrence. Accordingly, resource-based competitive exclusion should be rare. In contrast, contemporary ecosystem models typically predict an exclusion of larger phytoplankton size classes under low-nutrient conditions, an outcome interpreted as reflecting the competitive advantage of small cells having much higher nutrient ‘affinities’ than larger cells. Here, we develop mechanistically-focused expressions for steady-state, nutrient-limited phytoplankton growth that are consistent with the discrete, distantly-spaced cells of natural populations. These expressions, when encompassed in a phytoplankton-zooplankton model, yield sustained diversity across all size classes over the full range in nutrient concentrations observed in the ocean. In other words, our model does not exhibit resource-based competitive exclusion between size classes previously associated with size-dependent differences in nutrient ‘affinities’.

Nutrient limitation of phytoplankton in three tributaries of Chesapeake Bay: Detecting responses following nutrient reductions

Many coastal ecosystems suffer from eutrophication, algal blooms, and dead zones due to excessive anthropogenic inputs of nitrogen (N) and phosphorus (P). This has led to regional restoration efforts that focus on managing watershed loads of N and P. In Chesapeake Bay, the largest estuary in the United States, dual nutrient reductions of N and P have been pursued since the 1980s. However, it remains unclear whether nutrient limitation – an indicator of restriction of algal growth by supplies of N and P – has changed in the tributaries of Chesapeake Bay following decades of reduction efforts. Toward that end, we analyzed historical data from nutrient-addition bioassay experiments and data from the Chesapeake Bay long-term water-quality monitoring program for six stations in three tidal tributaries (i.e., Patuxent, Potomac, and Choptank Rivers). Classification and regression tree (CART) models were developed using concurrent collections of water-quality parameters for each bioassay monitoring location during 1990-2003, which satisfactorily predicted the bioassay-based measures of nutrient limitation (classification accuracy = 96%). Predictions from the CART models using water-quality monitoring data showed enhanced nutrient limitation over the period of 1985-2020 at four of the six stations, including the downstream station in each of these three tributaries. These results indicate detectable, long-term water-quality improvements in the tidal tributaries. Overall, this research provides a new analytical tool for detecting signs of ecosystem recovery following nutrient reductions. More broadly, the approach can be adapted to other waterbodies with long-term bioassays and water-quality data sets to detect ecosystem recovery.

Nutrient Control of Phytoplankton Abundance and Biomass, and Microplankton Assemblage Structure in the Lower Columbia River (Vancouver, Washington, USA)

Nutrient limitation of phytoplankton is common but by no means universal in large temperate rivers. Previous field studies in the Columbia River, USA, are suggestive of nutrient limitations of phytoplankton, especially during summer, but this has never been tested experimentally. We therefore undertook monthly 5-day nutrient amendment incubation experiments from May–September 2018 using Columbia River water collected at Vancouver, Washington, USA. We compared replicate treatment bottles containing natural microplankton assemblages and amended nutrients (NO3, PO4 and SiO4 in combination) with replicate control bottles containing natural microplankton assemblages and ambient nutrients. Phytoplankton abundance and biomass were compared between treatments and controls on each day of each experiment, and microplankton assemblage structure was evaluated using Permutational Multivariate Analysis of Variance and Non-Metric Multi-Dimensional Scaling ordination on Day 0 (ambient) and Day 5 of each experiment. Nutrient amendment significantly affected phytoplankton abundance and biomass, particularly in June–August, although this varied between taxa (e.g., cyanobacteria, dinoflagellates, flagellates and ciliates showed more frequent positive responses than chlorophytes and diatoms did). Abundance-based microplankton assemblage structure was significantly correlated with PO4, SiO4 and NO3 concentrations, and BIOENV procedure in R revealed that the best subset of explanatory variables included SiO4 and NO3 concentrations. Biomass-based assemblage structure was significantly correlated with SiO4 and NO3, although BIOENV explanatory variables included only SiO4. These results are suggestive of summertime nutrient control of phytoplankton abundance and biomass, as well as microplankton composition, in the lower Columbia River, at least during some months. Since eutrophication is increasing in the watershed, this could have important implications for higher level consumers (e.g., zooplankton and out-migrating juvenile salmon).

Environmental Regulation of Photosynthetically Produced Dissolved Organic Carbon by Phytoplankton Along a Subtropical Estuarine Bay

Photosynthetically produced dissolved organic carbon (PDOC) released by marine phytoplankton has an important significance on the marine carbon cycle and on the growth of heterotopic bacteria (HB). However, PDOC is often neglected in the estimation of global marine primary productivity and carbon sequestration capacity. This work studied the issue of PDOC and its environmental regulation mechanism through field investigation and lab experiments in an estuarine bay during southwest and northeast monsoon. We used the percentage of extracellular release (PER) as a key indicator to evaluate the contribution of PDOC to total primary productivity (TPP). We also compared PER among different seasons and sectors and then analyzed the inter-relationship between PDOC and bacterial carbon demand (BCD), size-fractionated phytoplankton, bacterial production (BP), and TPP. We finally discussed the impact factors of PDOC productivity. The results showed that the average contribution of PDOC to TPP in Qinzhou bay could reach 15% during two seasons, which satisfies about 25% of the carbon requirement by HB in the bay. Multiple factors contribute to the seasonal (SW monsoon 13% < NE monsoon 18%) and sectoral variation (outer bay 26% > middle bay 17% > inner bay 7%) in PER, the most significant of which are salinity and nitrogen to phosphorus ratios. PER is also related to phytoplankton community structure and nutrient limitation, the higher PER in the outer bay is attributed to the dual effect of picophytoplankton being the predominant species and a severe imbalance in the nitrogen to phosphorus ratio, both of which lead to increased phytoplankton PDOC release. On the other hand, the lower PER in the inner bay was mainly due to the dominance of microphytoplankton and nitrogen to phosphorus ratio which is close to the redfield ratio. There is still a non-uniform conclusion on the relationship between PER and TPP, suggesting that this relationship needs to be evaluated not only in a horizontal-independent way but also through seasonal and vertical dimensions. The non-linear relationship between PDOC and BCD found in this study potentially indicated weak coupling between phytoplankton exudation and bacterial metabolism. In addition to phytoplankton exudate, HB is still dependent on other sources of DOC to meet their carbon requirements.

Winter Time Nutrient Limitation of Phytoplankton in North Carolina Water Supply Reservoirs and An Inflowing Stream

Abstract In aquatic ecosystems, phytoplankton growth may be limited by one or more essential nutrients, a phenomenon known as nutrient limitation. Nutrient limitation can vary spatially, including within and among systems, and temporally, including across seasons and years. For systems that are susceptible to eutrophication (the nutrient over-enrichment of water bodies), understanding patterns of nutrient limitation can be useful for water resource managers. We conducted phytoplankton growth bioassays in late February 2019 to analyze the nutrient limitation status of two water supply reservoirs (Oak Hollow Lake and City Lake), as well as an inflowing stream (Boulding Branch), in the North Carolina Piedmont. Two sites within each reservoir were assessed, including one site close to the water supply intake and one site further upstream. Each laboratory bioassay consisted of three nutrient addition treatments: nitrogen (N+), phosphorus (P+), and nitrogen and phosphorus (NP+), as well as a control. After 48 hr incubation under a growth light, chlorophyll a was measured in all samples, including initial samples taken prior to growth light incubation, and ANOVA was used to determine significant differences among treatments. The City Lake upstream site was P-limited, whereas the site closer to the intake was co-limited by N and P. The Oak Hollow downstream site closest to the water supply intake was P-limited while the Oak Hollow upstream site was not nutrient limited. Similarly, Boulding Branch was not nutrient limited. These results illustrate the spatial complexity of nutrient limitation. Further studies are needed to better understand this complexity, as well as examine seasonal and interannual patterns of nutrient limitation in these critical water supply reservoirs.

Varying influence of phytoplankton biodiversity and stoichiometric plasticity on bulk particulate stoichiometry across ocean basins

Concentrations and elemental ratios of suspended particulate organic matter influence many biogeochemical processes in the ocean, including patterns of phytoplankton nutrient limitation and links between carbon, nitrogen and phosphorus cycles. Here we present direct measurements of cellular nutrient content and stoichiometric ratios for discrete phytoplankton populations spanning broad environmental conditions across several ocean basins. Median cellular carbon-to-phosphorus and nitrogen-to-phosphorus ratios were positively correlated with vertical nitrate-to-phosphate flux for all phytoplankton groups and were consistently higher for cyanobacteria than eukaryotes. Light and temperature were inconsistent predictors of stoichiometric ratios. Across nutrient-rich and phosphorus-stressed biomes in the North Atlantic, but not in the nitrogen-stressed tropical North Pacific, we find that a combination of taxonomic composition and environmental acclimation best predict bulk particulate organic matter composition. Our findings demonstrate the central role of plankton biodiversity and plasticity in controlling linkages between ocean nutrient and carbon cycles in some regions.

Carbon fixation by the phytoplankton community across Lake Winnipeg

Lakes are important sites for carbon fixation and carbon dioxide (CO2) exchange with the atmosphere. Carbon fixation rates have not previously been published for Lake Winnipeg but are important for quantifying the lake’s role in the regional greenhouse gas budget and the lake’s trophic structure and fish habitat. This study measured net ecosystem production (NEP), gross primary production (GPP), and gross respiration (GR) across the lake using a custom-built automated incubator connected to a ship’s water intake during a research cruise between July 31 and August 17, 2018 on Lake Winnipeg. The incubator estimated NEP, GR, and GPP every 60 min while moving along the ship’s track and at anchor, providing high-resolution data that are not obtainable through conventional incubations. The mean NEP for Lake Winnipeg during our survey was −8.4 ± 5.6 g C m−2 d−1, suggesting that the lake was net heterotrophic and thus a net CO2 source to the atmosphere during the 2018 summer cruise. The high-resolution data revealed significant spatiotemporal variability, including short-lived, highly net productive events that preceded remotely sensed chlorophyll a blooms by several days. Conversely, in regions with high chlorophyll a concentrations, we observed strong net heterotrophy and low nutrients, suggesting respiration was fueled by the degradation of mature, nutrient-limited phytoplankton blooms. The incubator system used in this study demonstrated its utility for monitoring rapid changes in NEP over short spatial scales in a lake which shows heightened regional variability in its physical, biogeochemical, and biological make-up.

Chemical contamination alters the interactions between bacteria and phytoplankton

Bacteria and phytoplankton are key players in aquatic ecosystem functioning. Their interactions mediate carbon transfer through the trophic web. Chemical contamination can alter the function and diversity of phytoplankton and bacterioplankton, with important consequences for ecosystem functioning. The aim of the present study was to assess the impact of chemical contamination on the interactions between both biological compartments. Two contrasting marine coastal ecosystems, offshore waters and lagoon waters, were exposed to chemical contamination (artificial or produced from resuspension of contaminated sediment) in microcosms in four seasons characterized by distinct phytoplankton communities. Offshore waters were characterized by a complex phytoplankton–bacterioplankton network with a predominance of positive interactions between both compartments, especially with Haptophyta, Cryptophyta, and dinoflagellates. In contrast, for lagoon waters, the phytoplankton–bacterioplankton network was simpler with a prevalence of negative interactions with Ochrophyta, Cryptophyta, and flagellates. Contamination with an artificial mix of pesticides and trace metal elements resulted in a decrease in the number of interactions between phytoplankton and bacterioplankton, especially for offshore waters. Resuspension of contaminated sediment also altered the interactions between both compartments. The release of nutrients stored in the sediment allowed the growth of nutrient limited phytoplankton species with marked consequences for the interactions with bacterioplankton, with a predominance of positive interactions, whereas in lagoon waters, negative interactions were mostly observed. Overall, this study showed that chemical contamination and sediment resuspension resulted in significant effects on phytoplankton–bacterioplankton interactions that can alter the functioning of anthropogenic coastal ecosystems.

Nutrient limitation of phytoplankton in Chesapeake Bay: Development of an empirical approach for water-quality management

Understanding the temporal and spatial roles of nutrient limitation on phytoplankton growth is necessary for developing successful management strategies. Chesapeake Bay has well-documented seasonal and spatial variations in nutrient limitation, but it remains unknown whether these patterns of nutrient limitation have changed in response to nutrient management efforts. We analyzed historical data from nutrient bioassay experiments (1992–2002) and data from long-term, fixed-site water-quality monitoring program (1990–2017) to develop empirical approaches for predicting nutrient limitation in the surface waters of the mainstem Bay. Results from classification and regression trees (CART) matched the seasonal and spatial patterns of bioassay-based nutrient limitation in the 1992–2002 period much better than two simpler, non-statistical approaches. An ensemble approach of three selected CART models satisfactorily reproduced the bioassay-based results (classification rate = 99%). This empirical approach can be used to characterize nutrient limitation from long-term water-quality monitoring data on much broader geographic and temporal scales than would be feasible using bioassays, providing a new tool for informing water-quality management. Results from our application of the approach to 21 tidal monitoring stations for the period of 2007–2017 showed modest changes in nutrient limitation patterns, with expanded areas of nitrogen-limitation and contracted areas of nutrient saturation (i.e., not limited by nitrogen or phosphorus). These changes imply that long-term reductions in nitrogen load have led to expanded areas with nutrient-limited phytoplankton growth in the Bay, reflecting long-term water-quality improvements in the context of nutrient enrichment. However, nutrient limitation patterns remain unchanged in the majority of the mainstem, suggesting that nutrient loads should be further reduced to achieve a less nutrient-saturated ecosystem.

Drivers of short-term variability in phytoplankton production in an embayment of the southern Benguela upwelling system

St Helena Bay (SHB), a retention zone in the southern Benguela upwelling system, supports 40–50% of the region's primary productivity. It also experiences harmful algal blooms (HABs) and hypoxic conditions that are difficult to predict given the high sub-seasonal variability. To better understand this variability, net primary production (NPP), nitrate and ammonium uptake, and phytoplankton community composition were monitored for ten days in SHB during summer, the season of enhanced upwelling. A period of active upwelling (days 1–5) was followed by one of relaxation (days 6–10). During upwelling, the mixed layer was deeper than the euphotic zone and phytoplankton were light-limited, evidenced by high ambient nitrate concentrations and low rates of NPP and nitrate uptake. During relaxation, stratification increased, restricting phytoplankton production to a shallow euphotic zone in which nitrate was exhausted after three days; the subsequent decline in NPP and nitrate uptake rates confirm that nutrient availability rapidly succeeded light as the dominant control on productivity. Nanophytoplankton (mainly the small diatoms, Chaetoceros spp. and Skeletonema costatum) dominated the biomass, NPP, and nitrate uptake throughout the experiment. We attribute this to their ability to respond quickly to newly-upwelled nitrate and to sustain elevated nitrate uptake rates for longer than pico- and microphytoplankton. They may also engage in luxury nitrate uptake, storing nitrate intracellularly under high-nutrient conditions and assimilating it later when nutrients are depleted. Additionally, Chaetoceros spp. and S. costatum are chain-formers, benefitting from the advantages of being small (i.e., as individual cells) and large (i.e., when aggregated into chains). A weakening of surface stratification late in the experiment may have prevented dinoflagellates, some of which are HAB species, from succeeding the diatoms. One implication of this is that understanding the rapid cycling between light and nutrient limitation of SHB phytoplankton, induced by an actively-upwelling versus stratified water column, may improve our capacity to anticipate HABs and the associated hypoxic events.

Concordance of chemically inferred and assayed nutrient limitation of phytoplankton along a depth gradient of alpine lakes in the Canadian Rockies

Deposition of nutrients over mountainous regions in many parts of the world is increasing due to human activities. To assess the level of agreement between chemically-inferred and experimental evidence of the potential sensitivity of alpine lake ecosystems along the Eastern Front range of the Canadian Rockies to increased deposition of nitrogen (N) and phosphorus (P), nutrient limitation was investigated in 29 sites using water chemistry and nutrient amendment bioassays of phytoplankton. The two lines of evidence agreed in 55% of all cases with discrepancies mainly involving instances of co-limitation. Dissolved inorganic N:total P (DIN:TP) mass ratios frequently suggested P limitation while the corresponding bioassays supported meta-analytical evidence of the prevalence of co-limitation of freshwater primary producers. Based on previous evidence of low DIN:TP mass ratios and experimental N amendments stimulating algal communities in shallower alpine sites, we hypothesized that nutrient limitation of phytoplankton shifts from being P- to N-driven with decreasing lake depth. However, we found little evidence of N limitation from bioassays of phytoplankton from the shallower (< 2-m deep) lakes. Most phytoplankton were co-limited by P plus N amendments (42%), while 38% responded to only P, 10% to only N, and 10% to neither nutrient. Overall, DIN:TP mass ratios and bioassay results showed that chromophytes and chlorophytes drove the greater responsiveness of whole communities to P relative to N. Since the short-term bioassay experiments often demonstrated the alpine phytoplankton to be NP-co-limited, our findings highlight a need for further research into potential synergistic fertilizing effects of anthropogenic N emissions and P deposition from forest fires on lake ecosystems at high elevations.

Decline in Atlantic Primary Production Accelerated by Greenland Ice Sheet Melt

AbstractProjections of climate impacts on marine net primary production (NPP) are reliant on Earth System Models (ESMs) that do not contain dynamic ice sheets. We assess the impact of potential Greenland ice sheet meltwater on projections of 21st century NPP using idealized ESM simulations. Under an extreme melt scenario, corresponding to 21st century sea level rise close to 2 m, Greenland meltwater amplified the decline in global NPP from a decrease of 3.2 PgC/yr to a decrease of 4.5 PgC/yr, relative to present. This additional reduction in NPP predominately occurs in the North Atlantic subtropical and subpolar gyres, as well as Atlantic eastern boundary upwelling systems. Accelerated NPP declines are the result of both surface freshening and reductions in upwelling‐favorable winds enhancing phytoplankton nutrient limitation. Our findings indicate that including a dynamic Greenland ice sheet in ESMs could have large impacts on projections of future ocean circulation and biogeochemistry.

Relevance of Nutrient-Limited Phytoplankton Production and Its Bacterial Remineralization for Carbon and Oxygen Fluxes in the Baltic Sea

The Baltic Sea is prone to oxygen deficiency due to the restricted water exchange with the North Sea in coincidence with a high biological oxygen demand. The partitioning of organic carbon between respiration, accumulation and export is co-determined by phytoplankton primary production and its subsequent bacterial remineralization. Here, we investigated net phytoplankton primary production, heterotrophic bacterial biomass production and dark CO2 fixation by on-board incubations with radiolabeled tracers in the Baltic Proper and in the Gulf of Riga after the main spring bloom. Results show that low phytoplankton standing stocks of ≤1.6 µg chlorphyll a L-1 sustained net primary production of 161-724 mg C m-2 d-1 under nitrogen limitation. Estimates of bacterial carbon remineralization suggest that freshly produced organic carbon was supplied to the aphotic zone at all stations. In the southern Baltic Proper, net primary production exceeded the bacterial carbon demand in the upper mixed layer, suggesting that organic matter derived from nutrient-limited primary production was available for export to bacterial communities below the oxycline. On average, 46% of heterotrophic bacterial production was mediated in oxygen minimum zones, revealing the high importance of organic matter recycling under hypoxic and anoxic conditions for the carbon budget. Dark CO2 fixation of up to 4.33 µg C L-1 d-1 in sulfide-free waters equaled 9-54% of the co-inciding heterotrophic bacterial carbon demand and may have provided another organic carbon source for heterotrophic activity. Substantially higher dark CO2 fixation up to 25.46 µg C L-1 d-1 was determined in sulfidic waters. Since our study was conducted five months after the major Baltic inflow event in winter 2014/2015, potential effects of deep water ventilation could be investigated. In the Gotland Basin, heterotrophic bacterial production in renewed oxygen-rich bottom water was similar to that in the uplifted oxygen-deficient former bottom water, while it was significantly reduced in sulfidic waters. Hence, our results suggest that the removal of hydrogen sufide by inflow events has a high potential to increase bacterial carbon remineralization.