Search

Billions of years ago, the Earth's waters were dominated by cyanobacteria. These microbes amassed to such formidable numbers, they ushered in a new era—starting with the Great Oxidation Event—fuelled by oxygenic photosynthesis. Throughout the following eon, cyanobacteria ceded portions of their global aerobic power to new photoautotrophs with the rise of eukaryotes (i.e. algae and higher plants), which co-existed with cyanobacteria in aquatic ecosystems. Yet while cyanobacteria's ecological success story is one of the most notorious within our planet's biogeochemical history, scientists to this day still seek to unlock the secrets of their triumph. Now, the Anthropocene has ushered in a new era fuelled by excessive nutrient inputs and greenhouse gas emissions, which are again reshaping the Earth's biomes. In response, we are experiencing an increase in global cyanobacterial bloom distribution, duration, and frequency, leading to unbalanced, and in many instances degraded, ecosystems. A critical component of the cyanobacterial resurgence is the freshwater-marine continuum: which serves to transport blooms, and the toxins they produce, on the premise that “water flows downhill”. Here, we identify drivers contributing to the cyanobacterial comeback and discuss future implications in the context of environmental and human health along the aquatic continuum. This Minireview addresses the overlooked problem of the freshwater to marine continuum and the effects of nutrients and toxic cyanobacterial blooms moving along these waters. Marine and freshwater research have historically been conducted in isolation and independently of one another. Yet, this approach fails to account for the interchangeable transit of nutrients and biology through and between these freshwater and marine systems, a phenomenon that is becoming a major problem around the globe. This Minireview highlights what we know and the challenges that lie ahead.

Estuaries function as important transporters, transformers, and producers of organic matter (OM). Along the freshwater to saltwater gradient, the composition of OM is influenced by physical and biogeochemical processes that change spatially and temporally, making it difficult to constrain OM in these ecosystems. In addition, many of the environmental parameters (temperature, precipitation, riverine discharge) controlling OM are expected to change due to climate change. To better understand the environmental drivers of OM quantity (concentration) and quality (absorbance, fluorescence), we assessed both dissolved OM (DOM) and particulate OM (POM) spatially, along the freshwater to saltwater gradient and temporally, for a full year. We found seasonal differences in salinity throughout the estuary due to elevated riverine discharge during the late fall to early spring, with corresponding changes to OM quantity and quality. Using redundancy analysis, we found DOM covaried with salinity (adjusted r2 = 0.35, 0.41 for surface and bottom), indicating terrestrial sources of DOM in riverine discharge were the dominant DOM sources throughout the estuary, while POM covaried with environmental indictors of terrestrial sources (turbidity, adjusted r2 = 0.16, 0.23 for surface and bottom) as well as phytoplankton biomass (chlorophyll-a, adjusted r2 = 0.25, 0.14 for surface and bottom). Responses in OM quantity and quality observed during the period of elevated discharge were similar to studies assessing OM quality following extreme storm events suggesting that regional changes in precipitation, as predicted by climate change, will be as important in changing the estuarine OM pool as episodic storm events in the future.

China has sought to address water pollution in the last decade by introducing a wide range of laws and regulations (Table S1), which led to nationwide water quality improvement. However, recent quantitative assessment of progress toward Sustainable Development Goals (SDGs) in China suggests that some SDGs underpinning goals for water pollution and biodiversity have not been met. Specifically, tests on water quality improvement in lakes have had contradictory results, leading to confusion about water quality improvement in China.

Estuaries regulate transport of dissolved organic carbon (DOC) from land to ocean. Export of terrestrial DOC from coastal watersheds is exacerbated by increasing major rainfall and storm events and human activities, leading to pulses of DOC that are shunted through rivers downstream to estuaries. Despite an upward trend of extreme events, the fate of the pulsed terrestrial DOC in estuaries remains unclear. We analyzed the effects of seven major tropical cyclones (TC) from 1999 to 2017 on the quantity and fate of DOC in the Neuse River Estuary (NC, USA). Significant TC-induced increases in DOC were observed throughout the estuary; the increase lasting from around 50 d at head-of-tide to over 6 months in lower estuary. Our results suggest that pulsed terrestrial DOC associated with TCs temporarily overwhelms the estuarine filter's abiotic and biotic degradation capacity under such high flow events, enhancing the shunt of terrestrial carbon to the coastal ocean.

The Defense Coastal/Estuarine Research Program (DCERP) was a 10-year multi-investigator project funded by the Department of Defense to improve understanding of ecosystem processes and their interactions with natural and anthropogenic stressors at the Marine Corps Base Camp Lejeune (MCBCL) located in coastal North Carolina. The project was aimed at facilitating ecosystem-based management (EBM) at the MCBCL and other coastal military installations. Because of its scope, interdisciplinary character, and duration, DCERP embodied many of the opportunities and challenges associated with EBM, including the need for explicit goals, system models, long-term perspectives, systems complexity, change inevitability, consideration of humans as ecosystem components, and program adaptability and accountability. We describe key elements of this program, its contributions to coastal EBM, and its relevance as an exemplar of EBM.

The Neuse River Estuary (North Carolina, USA) is a valuable ecosystem that has been affected by the expansion of agricultural and urban watershed activities over the last several decades. Eutrophication, as a consequence of enhanced anthropogenic nutrient loadings, has promoted high phytoplankton biomass, hypoxia, and fish kills. This study compares and contrasts three models to better understand how nutrient loading and other environmental factors control phytoplankton biomass, as chl-a, over time. The first model is purely statistical, while the second model mechanistically simulates both chl-a and nitrogen dynamics, and the third additionally simulates phosphorus. The models are calibrated to a multi-decadal dataset (1997–2018) within a Bayesian framework, which systematically incorporates prior information and accounts for uncertainties. All three models explain over one third of log-transformed chl-a variability, with the mechanistic models additionally explaining the majority of the variability in bioavailable nutrients (R2 > 0.5). By disentangling the influences of riverine nutrient concentrations, flows, and loadings on estuary productivity we find that concentration reductions, rather than total loading reductions, are the key to controlling estuary chl-a levels. The third model indicates that the estuary, even in its upstream portion, is rarely phosphorus limited, and will continue to be mostly nitrogen limited even under a 30% phosphorus reduction scenario. This model also predicts that a 10% change in nitrogen loading (flow held constant) will produce an approximate 4.3% change in estuary chl-a concentration, while the statistical model suggests a larger (10%) effect. Overall, by including a more detailed representation of environmental factors controlling algal growth, the mechanistic models generate chl-a forecasts with less uncertainty across a range of nutrient loading scenarios. Methodologically, this study advances the use of Bayesian methods for modeling the eutrophication dynamics of an estuarine system over a multi-decadal period.

Climate warming in combination with nutrient enrichment can greatly promote phytoplankton proliferation and blooms in eutrophic waters. Lake Taihu, China, is a large, shallow and eutrophic system. Since 2007, this lake has experienced extensive nutrient input reductions aimed at controlling cyanobacterial blooms. However, intense cyanobacterial blooms have persisted through 2017 with a record-setting bloom occurring in May 2017. Causal analysis suggested that this bloom was sygenerically driven by high external loading from flooding in 2016 in the Taihu catchment and a notable warmer winter during 2016/2017. High precipitation during 2016 was associated with a strong 2015/2016 El Niño in combination with the joint effects of Atlantic Multi-decadal Oscillation (AMO) and Pacific Decadal Oscillation (PDO), while persistent warmth during 2016/2017 was strongly related to warm phases of AMO and PDO. The 2017 blooms elevated water column pH and led to dissolved oxygen depletion near the sediment, both of which mobilized phosphorus from the sediment to overlying water, further promoting cyanobacterial blooms. Our finding indicates that regional climate anomalies exacerbated eutrophication via a positive feedback mechanism, by intensifying internal nutrient cycling and aggravating cyanobacterial blooms. In light of global expansion of eutrophication and blooms, especially in large, shallow and eutrophic lakes, these regional effects of climate anomalies are nested within larger scale global warming predicted to continue in the foreseeable future.

Hypoxia is common feature of eutrophic estuaries and semi-enclosed seas globally. One of the key factors driving hypoxia is nitrogen pollution. To gain more insight into the effects of hypoxia on estuarine nitrogen cycling, we measured potential nitrate reduction rates at different salinities and levels of hypoxia in a eutrophic temperate microtidal estuary, the Neuse River Estuary, North Carolina, USA. We also tested the effect of hydrogen sulfide and ferrous iron additions on the nitrate reduction pathways. Overall, DNRA dominated over denitrification in this periodically hypoxic estuary and there was no correlation between the potential nitrate reduction rates, salinity, or dissolved oxygen. However, when hypoxia lasted several months, denitrification capacity was almost completely lost, and nearly all nitrate added to the sediment was reduced via DNRA. Additions of hydrogen sulfide stimulated DNRA over denitrification. Additions of ferrous iron stimulated nitrate consumption; however, the end product of nitrate consumption was not clear. Interestingly, substantial nitrous oxide formation occurred in sediments that had experienced prolonged hypoxia and were amended with nitrate. Given expanding hypoxia predicted with climate change scenarios and the increasing nitrate loads to coastal systems, coastal sediments may lose their capability to mitigate nitrogen pollution due to DNRA dominating over denitrification during extended hypoxic periods.

The global expansion of harmful cyanobacterial blooms (CyanoHABs) poses an increasing threat to public health. CyanoHABs are characterized by the production of toxic metabolites known as cyanotoxins. Human exposure to cyanotoxins is challenging to forecast, and perhaps the least understood exposure route is via inhalation. While the aerosolization of toxins from marine harmful algal blooms (HABs) has been well documented, the aerosolization of cyanotoxins in freshwater systems remains understudied. In recent years, spray aerosol (SA) produced in the airshed of the Laurentian Great Lakes (United States and Canada) has been characterized, suggesting that freshwater systems may impact atmospheric aerosol loading more than previously understood. Therefore, further investigation regarding the impact of CyanoHABs on human respiratory health is warranted. This review examines current research on the incorporation of cyanobacterial cells and cyanotoxins into SA of aquatic ecosystems which experience HABs. We present an overview of cyanotoxin fate in the environment, biological incorporation into SA, existing data on cyanotoxins in SA, relevant collection methods, and adverse health outcomes associated with cyanotoxin inhalation.

Cyanobacterial harmful algal bloom (CyanoHAB) proliferation is a global problem impacting ecosystem and human health. Western Lake Erie (WLE) typically endures two highly toxic CyanoHABs during summer: a Microcystis spp. bloom in Maumee Bay that extends throughout the western basin, and a Planktothrix spp. bloom in Sandusky Bay. Recently, the USA and Canada agreed to a 40% phosphorus (P) load reduction to lessen the severity of the WLE blooms. To investigate phosphorus and nitrogen (N) limitation of biomass and toxin production in WLE CyanoHABs, we conducted in situ nutrient addition and 40% dilution microcosm bioassays in June and August 2019. During the June Sandusky Bay bloom, biomass production as well as hepatotoxic microcystin and neurotoxic anatoxin production were N and P co-limited with microcystin production becoming nutrient deplete under 40% dilution. During August, the Maumee Bay bloom produced microcystin under nutrient repletion with slight induced P limitation under 40% dilution, and the Sandusky Bay bloom produced anatoxin under N limitation in both dilution treatments. The results demonstrate the importance of nutrient limitation effects on microcystin and anatoxin production. To properly combat cyanotoxin and cyanobacterial biomass production in WLE, both N and P reduction efforts should be implemented in its watershed.

Harmful cyanobacterial blooms (CyanoHABs) are a rapidly proliferating global problem, threatening the use and sustainability of our freshwater resources. In recent decades, the United States, China, and other developed and developing countries threatened by CyanoHAB expansion have established collaborative efforts aimed at mitigating and managing this environmental and human health problem. However, an escalating negative political climate and restrictive policies on scientific exchange threaten these efforts. In this Perspective, I point to progress that has been made to counter the CyanoHAB problem on U.S.–Chinese fronts through our collaborations, which have been mutually beneficial from research and academic perspectives. Much like global efforts now needed to control pandemics, we are all “in the same boat” when to comes to countering the threat CyanoHABs pose for drinkable, swimmable, and fishable freshwater supplies and human health.

Bacteria play key roles in the function and diversity of aquatic systems, but aside from study of specific bloom systems, little is known about the diversity or biogeography of bacteria associated with harmful cyanobacterial blooms (cyanoHABs). CyanoHAB species are known to shape bacterial community composition and to rely on functions provided by the associated bacteria, leading to the hypothesized cyanoHAB interactome, a coevolved community of synergistic and interacting bacteria species, each necessary for the success of the others. Here, we surveyed the microbiome associated with Microcystis aeruginosa during blooms in 12 lakes spanning four continents as an initial test of the hypothesized Microcystis interactome. We predicted that microbiome composition and functional potential would be similar across blooms globally. Our results, as revealed by 16S rRNA sequence similarity, indicate that M. aeruginosa is cosmopolitan in lakes across a 280° longitudinal and 90° latitudinal gradient. The microbiome communities were represented by a wide range of operational taxonomic units and relative abundances. Highly abundant taxa were more related and shared across most sites and did not vary with geographic distance, thus, like Microcystis, revealing no evidence for dispersal limitation. High phylogenetic relatedness, both within and across lakes, indicates that microbiome bacteria with similar functional potential were associated with all blooms. While Microcystis and the microbiome bacteria shared many genes, whole-community metagenomic analysis revealed a suite of biochemical pathways that could be considered complementary. Our results demonstrate a high degree of similarity across global Microcystis blooms, thereby providing initial support for the hypothesized Microcystis interactome.

Cyanobacterial harmful algal blooms (CyanoHABs) are an increasingly common feature of large, eutrophic lakes. Non-N2-fixing CyanoHABs (e.g., Microcystis) appear to be proliferating relative to N2-fixing CyanoHABs in systems receiving increasing nutrient loads. This shift reflects increasing external nitrogen (N) inputs, and a > 50-year legacy of excessive phosphorus (P) and N loading. Phosphorus is effectively retained in legacy-impacted systems, while N may be retained or lost to the atmosphere in gaseous forms (e.g., N2, NH3, N2O). Biological control on N inputs versus outputs, or the balance between N2 fixation versus denitrification, favors the latter, especially in lakes undergoing accelerating eutrophication, although denitrification removal efficiency is inhibited by increasing external N loads. Phytoplankton in eutrophic lakes have become more responsive to N inputs relative to P, despite sustained increases in N loading. From a nutrient management perspective, this suggests a need to change the freshwater nutrient limitation and input reduction paradigms; a shift from an exclusive focus on P limitation to a dual N and P co-limitation and management strategy. The recent proliferation of toxic non-N2-fixing CyanoHABs, and ever-increasing N and P legacy stores, argues for such a strategy if we are to mitigate eutrophication and CyanoHAB expansion globally.

Picophytoplankton (PicoP) are increasingly recognized as significant contributors to primary productivity and phytoplankton biomass in coastal and estuarine systems. Remarkably though, PicoP composition is unknown or not well-resolved in several large estuaries including the semi-lagoonal Neuse River Estuary (NRE), a tributary of the second largest estuary-system in the lower USA, the Pamlico-Albemarle Sound. The NRE is impacted by extreme weather events, including recent increases in precipitation and flooding associated with tropical cyclones. Here we examined the impacts of moderate to extreme (Hurricane Florence, September 2018) precipitation events on NRE PicoP abundances and composition using flow cytometry, over a 1.5 year period. Phycocyanin-rich Synechococcus-like cells were the most dominant PicoP, reaching ~ 106 cells mL−1, which highlights their importance as key primary producers in this relatively long residence-time estuary. Ephemeral “blooms” of picoeukaryotic phytoplankton (PEUK) during spring and after spikes in river flow were also detected, making PEUK periodically major contributors to PicoP biomass (up to ~ 80%). About half of the variation in PicoP abundance was explained by measured environmental variables. Temperature explained the most variation (24.5%). Change in total dissolved nitrogen concentration, an indication of increased river discharge, explained the second-most variation in PicoP abundance (15.9%). The short-term impacts of extreme river discharge from Hurricane Florence were particularly evident as PicoP biomass was reduced by ~ 100-fold for more than 2 weeks. We conclude that precipitation is a highly influential factor on estuarine PicoP biomass and composition, and show how ‘wetter’ future climate conditions will have ecosystem impacts down to the smallest of phytoplankton.

Managing and mitigating the global expansion of toxic cyanobacterial harmful algal blooms (CyanoHABs) is a major challenge facing researchers and water resource managers. Various approaches, including nutrient load reduction, artificial mixing and flushing, omnivorous fish removal, algaecide applications and sediment dredging, have been used to reduce bloom occurrences. However, managers now face the additional challenge of having to address the effects of climate change on watershed hydrological and nutrient load dynamics, water temperature, mixing regime and internal nutrient cycling. Rising temperatures and increasing frequencies and magnitudes of extreme weather events, including tropical cyclones, extratropical storms, floods and droughts, all promote CyanoHABs and affect the efficacy of ecosystem remediation measures. These climatic changes will likely require setting stricter nutrient (including both nitrogen and phosphorus) reduction targets for bloom control in affected waters. In addition, the efficacy of currently used methods to reduce CyanoHABs will need to be re-evaluated in light of the synergistic effects of climate change with nutrient enrichment.

Estuarine-coastal ecosystems are rich areas of the global ocean with elevated rates of organic matter production supporting major fisheries. Net and gross primary production (NPP, GPP) are essential properties of these ecosystems, characterized by high spatial, seasonal, and inter-annual variability associated with climatic effects on hydrology. Over 20 years ago, Nixon defined the trophic classification of marine ecosystems based on annual phytoplankton primary production (APPP), with categories ranging from “oligotrophic” to “hypertrophic”. Source data consisting of shipboard measurements of NPP and GPP from 1982 to 2004 for Chesapeake Bay in the mid-Atlantic region of the United States supported estimates of APPP from 300 to 500 g C m−2 yr−1, corresponding to “eutrophic” to “hypertrophic” categories. Here, we developed generalized additive models (GAM) to interpolate the limited spatio-temporal resolution of source data. Principal goals were: (1) to develop predictive models of NPP and GPP calibrated to source data (1982 to 2004); (2) to apply the models to historical (1960s, 1970s) and monitoring (1985 to 2015) data with adjustments for nutrient loadings and climatic effects; (3) to estimate APPP from model predictions of NPP; (4) to test effects of simulated reductions of phytoplankton biomass or nutrient loadings on trophic classification based on APPP. Simulated 40% decreases of euphotic-layer chl-a or TN and NO2 + NO3 loadings led to decreasing APPP sufficient to change trophic classification from “eutrophic’ to “mesotrophic” for oligohaline (OH) and polyhaline (PH) salinity zones, and from “hypertrophic” to “eutrophic” for the mesohaline (MH) salinity zone of the bay. These findings show that improved water quality is attainable with sustained reversal of nutrient over-enrichment sufficient to decrease phytoplankton biomass and APPP.

Cyanobacterial harmful algal blooms (CyanoHABs) are a major threat to human and environmental health. As global proliferation of CyanoHABs continues to increase in prevalence, intensity, and toxicity, it is important to identify and integrate the underlying causes and controls of blooms in order to develop effective short- and long-term mitigation strategies. Clearly, nutrient input reductions should receive high priority. Legacy effects of multi-decadal anthropogenic eutrophication have altered limnetic systems such that there has been a shift from exclusive phosphorus (P) limitation to nitrogen (N) limitation and N and P co-limitation. Additionally, climate change is driving CyanoHAB proliferation through increasing global temperatures and altered precipitation patterns, including more extreme rainfall events and protracted droughts. These scenarios have led to the “perfect storm scenario”; increases in pulsed nutrient loading events, followed by persistent low-flow, long water residence times, favoring bloom formation and proliferation. To meet the CyanoHAB mitigation challenge, we must: (1) Formulate watershed and airshed-specific N and P input reductions on a sliding scale to meet anthropogenic and climatic forcings. (2) Develop CyanoHAB management strategies that incorporate current and anticipated climatic changes and extremes. (3) Make nutrient management strategies compatible with other physical-chemical-biological mitigation approaches, such as altering freshwater flow and flushing, dredging, chemical applications, introduction of selective grazers, etc. (4) Target CyanoHAB toxin production and developing management approaches to reduce toxin production. (5) Develop broadly applicable long-term strategies that incorporate the above recommendations.

Phosphorus (P) is a vital nutrient for algal growth. Aside from soluble reactive P (SRP), organic P (OP) is used by algae via alkaline phosphatase (AP) hydrolysis, which can play an important role in supplying P. Enzymatically-hydrolysable OP (EHP) can potentially be used as an indicator of bioavailability of P other than SRP in natural waters. We investigated the ecological significance of alkaline phosphatase activity (APA), EHP concentration and P turnover time in the inflowing rivers of Lake Taihu (Taihu) during three hydrologic periods. Results indicated high SRP concentration and low SRP demand by algal suppressed APA in the inflowing rivers, the highest proportion of OP mineralization rate (v) to the maximum reaction velocity of AP (Vmax) is only 14.7%. P turnover time of the inflowing rivers was generally from 3 to 7 days and in exceptional cases, it could exceed 10 days. The high EHP reserve and the sufficient AP for OP mineralization render the rivers a significant source of utilizable OP, further exacerbating eutrophication of Taihu.

Flooding from extreme weather events (EWE), such as hurricanes, exports large amounts of dissolved organic matter (DOM) to both estuaries and coastal waters globally. Hydrologic connectivity of wetlands to adjacent river channels during flood events can potentially have a major control on the DOM exported to coastal waters after EWEs. In this study, a geographic information system based flood model was used to: (1) determine the volume of flooded wetlands in a river corridor following Hurricane Matthew in 2016; (2) compute the resulting volume fluxes of DOM to the Neuse River Estuary-Pamlico Sound (NRE-PS), in eastern North Carolina and (3) use the flood model to quantify the wetland contribution to DOM export. The flood model-derived contributions were validated with a Bayesian Monte Carlo mixing model combining measurements of DOM quality: specific UV Absorbance at 254 nm (SUVA254), spectral slope ratio (SR), and stable isotope ratios of dissolved organic carbon (δ13C-DOC). Results indicated that (1) hydrologic connectivity of the freshwater riparian wetlands caused the wetlands to become the primary source of organic matter (OM) that was exported into the NRE-PS after Matthew and (2) this source lingered in these coastal waters in the months after the storm. Thus, in consideration of the pulse-shunt concept, EWE such as Hurricane Matthew cause pulses of DOM from wetlands, which were the primary source of the OM shunted from the terrestrial environment to the estuary and sound. Wetlands constituted ca. 48% of the annual loading of DOC into the NRE and 16% of DOC loading into the PS over a period of 30 days after Hurricane Matthew. Results were consistent with prior studies in this system, and other coastal ecosystems, that attributed a high reactivity of DOM as the underlying reason for large CO2 releases following EWE. Adapting the pulse-shunt concept to estuaries requires the addition of a “processing” step to account for the DOM to CO2 dynamics, thus a new pulse-shunt process is proposed to incorporate coastal waters. Our results suggest that with increasing frequency and intensity of EWE, strengthening of the lateral transfer of DOM from land to ocean will occur and has the potential to greatly impact coastal carbon cycling.

Globally, many shallow lakes have shifted from a clear macrophyte-dominated state to a turbid phytoplankton-dominated state due to eutrophication. Such shifts are often accompanied by toxic cyanobacterial blooms, with specialized traits including buoyancy regulation and nitrogen fixation. Previous work has focused on how these traits contribute to cyanobacterial competitiveness. Yet, little is known on how these traits affect the value of nutrient loading thresholds of shallow lakes. These thresholds are defined as the nutrient loading at which lakes shift water quality state. Here, we used a modelling approach to estimate the effects of traits on nutrient loading thresholds. We incorporated cyanobacterial traits in the process-based ecosystem model PCLake+, known for its ability to determine nutrient loading thresholds. Four scenarios were simulated, including cyanobacteria without traits, with buoyancy regulation, with nitrogen fixation, and with both traits. Nutrient loading thresholds were obtained under N-limited, P-limited, and colimited conditions. Results show that cyanobacterial traits can impede lake restoration actions aimed at removing cyanobacterial blooms via nutrient loading reduction. However, these traits hardly affect the nutrient loading thresholds for clear lakes experiencing eutrophication. Our results provide references for nutrient loading thresholds and draw attention to cyanobacterial traits during the remediation of eutrophic water bodies.

In 2016, Hurricane Matthew accounted for 25% of the annual riverine C loading to the Neuse River Estuary-Pamlico Sound, in eastern North Carolina. Unlike inland watersheds, dissolved organic carbon (DOC) was the dominant component of C flux from this coastal watershed and stable carbon isotope and chromophoric dissolved organic matter evidence indicated the estuary and sound were dominated by wetland-derived terrigenous organic matter sources for several months following the storm. Persistence of wetland-derived DOC enabled its degradation to carbon dioxide (CO2), which was supported by sea-to-air CO2 fluxes measured in the sound weeks after the storm. Under future increasingly extreme weather events such as Hurricane Matthew, and most recently Hurricane Florence (September 2018), degradation of terrestrial DOC in floodwaters could increase flux of CO2 from estuaries and coastal waters to the atmosphere.

The frequency and intensity of cyanobacterial blooms are increasing worldwide. Interactions between toxic cyanobacteria and aquatic microorganisms need to be critically evaluated to understand microbial drivers and modulators of the blooms. In this study, we applied 16S/18S rRNA gene sequencing and metabolomics analyses to measure the microbial community composition and metabolic responses of the cyanobacterium Microcystis aeruginosa in a coculture system receiving dissolved inorganic nitrogen and phosphorus (DIP) close to representative concentrations in Lake Taihu, China. M. aeruginosa secreted alkaline phosphatase using a DIP source produced by moribund and decaying microorganisms when the P source was insufficient. During this process, M. aeruginosa accumulated several intermediates in energy metabolism pathways to provide energy for sustained high growth rates and increased intracellular sugars to enhance its competitive capacity and ability to defend itself against microbial attack. It also produced a variety of toxic substances, including microcystins, to inhibit metabolite formation via energy metabolism pathways of aquatic microorganisms, leading to a negative effect on bacterial and eukaryotic microbial richness and diversity. Overall, compared with the monoculture system, the growth of M. aeruginosa was accelerated in coculture, while the growth of some cooccurring microorganisms was inhibited, with the diversity and richness of eukaryotic microorganisms being more negatively impacted than those of prokaryotic microorganisms. These findings provide valuable information for clarifying how M. aeruginosa can potentially modulate its associations with other microorganisms, with ramifications for its dominance in aquatic ecosystems.

Coastal North Carolina, USA, has experienced three extreme tropical cyclone-driven flood events since 1999, causing catastrophic human impacts from flooding and leading to major alterations of water quality, biogeochemistry, and ecological conditions. The apparent increased frequency and magnitudes of such events led us to question whether this is just coincidence or whether we are witnessing a regime shift in tropical cyclone flooding and associated ecosystem impacts. Examination of continuous rainfall records for coastal NC since 1898 reveals a period of unprecedentedly high precipitation since the late-1990’s, and a trend toward increasingly high precipitation associated with tropical cyclones over the last 120 years. We posit that this trend, which is consistent with observations elsewhere, represents a recent regime shift with major ramifications for hydrology, carbon and nutrient cycling, water and habitat quality and resourcefulness of Mid-Atlantic and possibly other USA coastal regions.

Long-term phytoplankton community changes indicate the trophic status under nutrient regulation conditions in eutrophic lakes, although the modulatory role of climate change scenarios in their indicative function has been underestimated. Hence, we analyzed the relative contributions of nutrient and climate factors to interannual and seasonal variations in the phytoplankton biomass and composition from 1992 to 2017 in Meiliang Bay, Lake Taihu, China. Discrete phytoplankton communities from five periods were classified according to their interannual features. Variations in the phytoplankton community composition were observed during the five periods, and these variations included a shift from exclusive cyanobacterial dominance before 2008 to diatom-cyanobacterial codominance in 2008–2010 and from cyanobacterial dominance in 2011–2014 to cyanobacterial-diatom codominance in 2015–2017. The phytoplankton biomass pattern typically peaked in summer, although peaks also occurred in winter 2008–2010 and autumn 2011–2014. Additionally, the phytoplankton biomass increased by threefold from 2015 to 2017, which might have been related to rising air temperatures and greater light availability. The variance in the phytoplankton community was significantly explained by nutrient (ammonium, nitrate and phosphate)and climatic (air temperature and wind speed)factors. However, the explaining effect of the factors varied among the five periods: nutrients strongly impacted the community composition from 1992 to 2007, whereas climatic variables became more important modulators after 2007. These results reveal that climatic factors play importance roles in shaping the phytoplankton community and cyanobacterial blooms and suggest that differences in the roles between specific climatic conditions should be considered. Future declines in cyanobacterial blooms require further dual nitrogen and phosphorus reduction and longer recovery times under current climate change scenarios in this and possibly other shallow eutrophic lakes.

With the expansion of urban, industry, and agriculture after World War II, eutrophication firstly emerged as a major water quality threat in small water bodies. As the increasing magnitudes and scales of nutrient pollution and habitat alteration, many of the world’s large lakes exhibit symptoms of eutrophication, e.g., toxic cyanobacterial blooms, deoxygenation, and habitat loss. These symptoms were noted in shallow large lakes such as Okeechobee and Winnebago (USA), Winnipeg (Canada), Peipsi (Estonia), Balaton (Hungry), Chaohu and Taihu (China), Kasumigaura (Japan); in shallow parts of large lakes including Lakes Champlain, Ontario, and Erie (Canada/USA), Huron (USA), Maracaibo (Venezuela), Victoria (Africa); and in segments of immense water bodies such as Tanganyika (Africa) (Table S1 online). These large aquatic ecosystems pose a tremendous challenge from mitigation and restoration perspectives.

A long-standing debate in ecology deals with the role of nitrogen and phosphorus in management and restoration of aquatic ecosystems. It has been argued that nutrient reduction strategies to combat blooms of phytoplankton or floating plants should solely focus on phosphorus (P). The underlying argument is that reducing nitrogen (N) inputs is ineffective because N 2 -fixing species will compensate for N deficits, thus perpetuating P limitation of primary production. A mechanistic understanding of this principle is, however, incomplete. Here, we use resource competition theory, a complex dynamic ecosystem model and a 32-year field data set on eutrophic, floating-plant dominated ecosystems to show that the growth of non-N 2 -fixing species can become N limited under high P and low N inputs, even in the presence of N 2 fixing species. N 2 -fixers typically require higher P concentrations than non-N 2 -fixers to persist. Hence, the N 2 fixers cannot deplete the P concentration enough for the non-N 2 -fixing community to become P limited because they would be outcompeted. These findings provide a testable mechanistic basis for the need to consider the reduction of both N and P inputs to most effectively restore nutrient over-enriched aquatic ecosystems.

Coincident climatic and human effects strongly influence water-quality properties in estuarine-coastal ecosystems around the world. Time-series data for a number of ecosystems reveal high spatio-temporal variability superimposed on secular trends traceable to nutrient over-enrichment. In this paper, we present new analyses of long-term data for Chesapeake Bay directed at several goals: (1) to distinguish trends from spatio-temporal variability imposed by climatic effects; (2) to assess long-term trends of water-quality properties reflecting degradation and recovery; (3) to propose numerical water-quality criteria as targets for restoration; (4) to assess progress toward attainment of these targets. The bay has experienced multiple impairments associated with nutrient over-enrichment since World War II, e.g., low dissolved oxygen (DO), decreased water clarity, and harmful algal blooms (HAB). Anthropogenic eutrophication has been expressed as increased chlorophyll-a (chl-a) driven by accelerated nutrient loading from 1945 to 1980. Management intervention led to decreased loading thereafter, but deleterious symptoms of excess nutrients persist. Climatic effects exemplified by irregular “dry” and “wet” periods in the last 30+ years largely explain high inter-annual variability of water-quality properties, requiring adjustments to resolve long-term trends. Here, we extend these analyses at a finer temporal scale to six decades of chl-a, Secchi depth, and nitrite plus nitrate (NO 2 + NO 3 ) data to support trend analyses and the development of numerical water-quality criteria. The proposed criteria build on a conceptual model emphasizing the need to distinguish climatic and human effects in gauging progress to reverse eutrophication in estuarine-coastal ecosystems.

As the interface between riverine and coastal systems, estuaries play a key role in receiving, transporting, and processing terrestrial organic carbon prior to export to downstream coastal systems. Estuaries can switch from terrestrial organic carbon reactors under low river flow to pipelines under high flow, but it remains unclear how estuarine terrestrial organic carbon processing responds to the full spectrum of discharge conditions, which are bracketed by these high and low discharge events. The amount of terrestrial dissolved organic carbon and colored dissolved organic matter imported, processed, and exported was assessed for riverine discharge events spanning from the 4th to 99th flow quantiles in the Neuse River Estuary, North Carolina, USA using spatially and temporally (July 2015–December 2016) resolved measurements. The extent of dissolved organic matter processing in the estuary under various flow conditions was estimated using a non-steady state box model to calculate estuary-wide terrestrial dissolved organic carbon and colored dissolved organic matter source & sink terms. Under mid-range riverine discharge conditions (4th to 89th flow quantiles), the Neuse River Estuary was a sink for terrestrial dissolved organic carbon, retaining and/or processing (i.e., flocculation; photochemical and microbial degradation) on average ∼29% of terrestrial dissolved organic carbon. Following floods due to extreme precipitation events (99th flow quantile), however, over 99% of the terrestrial dissolved organic carbon loaded from the riverine end-member was exported directly to the downstream coastal system. Following such extreme weather events, the estuary acts as a pipeline for direct export of terrestrial dissolved organic carbon, drastically altering the amount and quality of dissolved organic carbon loaded to downstream coastal systems. This has important implications under future climate scenarios, where extreme weather events are expected to increase.

The concept of lakes “evolving” phosphorus (P) limitation has persisted in limnology despite limited direct evidence. Here, we developed a simple model to broadly characterize nitrogen (N) surpluses and deficits, relative to P, in lakes and compared the magnitude of this imbalance to estimates of N gains and losses through biological N transformations. The model suggested that approximately half of oligotrophic lakes in the U.S.A. had a stoichiometric N deficit, but 72–89% of eutrophic and hypereutrophic lakes, respectively, had a similar N deficit. Although reactive N appeared to accumulate in the most oligotrophic lakes, net denitrification perpetuated the N deficit in more productive lakes. Productive lakes exported reactive N via biological N transformations regardless of their N deficit. The lack of N accumulation through N fixation underscores the need for a modern eutrophication management approach focused on reducing total external nutrient loads, including both N and P.

A severe cyanobacterial bloom in the mainstem of a large Chinese river was first reported from China. The Qiantang River is the longest river in the Zhejiang province, southeast China. It provides drinking water supply to ~ 16 million people, including Hangzhou city. Fifteen sites along the Qiantang River (including upper, middle (Fuchunjiang Reservoir), and lower reaches and tributaries) were sampled between August 13 and September 9, 2016 to conduct a preliminary examination of the outbreak of Microcystis blooms. Laboratory investigation revealed that Microcystis spp. are dominant in the Fuchunjiang Reservoir (an overflow reservoir on the mainstem of the Qiantang River) with an extremely high cell density of 2.3 × 108 cells/L, leading to a severe bloom in the mainstem of the Qiantang River. Investigations of the meteorological, hydrological, and nutrient characteristics associated with the bloom indicated that extremely dry (6.8 mm rainfall from August 13 to September 9, 2016) and hot (32 consecutive days of temperatures > 30 °C from July 20 to August 31, 2016) weather might be the key factors triggering the bloom. Additionally, the extremely low flow of the tributary, Lanjiang River (142 ± 56 m3/s from August 13 to September 9), and its high nutrient background, favored the bloom. While nutrient reductions are important, the most immediate and effective management approach might be to implement appropriate minimal flow conditions to mitigate the blooms.

Extreme climatic events like floods and hurricanes have the potential to significantly alter coastal carbon cycling. However, due to the challenges associated with sampling these events, they can be difficult to incorporate into regional and global carbon budgets. To address this data gap, we bracket a major flooding event associated with the passing of Hurricane Joaquin (October 2015) with direct and high-resolution pCO2 measurements in the Neuse (NeuseRE) and New River Estuary (NewRE), North Carolina. Enhanced river discharge quickly flushed the relatively small NewRE, causing residence time to fall from 90 to 9 days, while the larger NeuseRE responded relatively slowly to flooding. This period of rapid flushing coincided with a significant increase in CO2 fluxes. The effect of cooler flood-waters, which reduce pCO2, was counteracted by allochthonous DIC inputs, which drove large increases in pCO2 relative to dissolved O2. The spatial distribution of carbonate buffering differed between estuaries, enhancing CO2 fluxes in the NewRE (178 mmol C m−2 d−1), while partially limiting air-water exchange in the NeuseRE (62 mmol C m−2 d−1). While windy storms may drive larger CO2 fluxes from estuaries, we show that flooding events can also contribute significantly to annual carbon budgets. CO2 emissions during this ~14-day flood period accounted for 31% (NeuseRE) to 44% (NewRE) of the total annual CO2 flux. Our findings show that sufficient spatial and temporal coverage during storms is necessary for estuarine CO2 fluxes to be reliably assessed over annual or longer time scales.

Anthropogenic nutrient overenrichment, coupled with rising temperatures, and an increasing frequency of extreme hydrologic events (storms and droughts) are accelerating eutrophication and promoting the expansion of harmful algal blooms (HABs) across the freshwater-to-marine continuum. All HABs - with a focus here on cyanobacterial blooms - pose serious consequences for water supplies, fisheries, recreational uses, tourism, and property values. As nutrient loads grow in watersheds, they begin to compound the effects of legacy stores. This has led to a paradigm shift in our understanding of how nutrients control eutrophication and blooms. Phosphorus (P) reductions have been traditionally prescribed exclusively for freshwater systems, while nitrogen (N) reductions were mainly stressed for brackish and coastal waters. However, because most systems are hydrologically interconnected, single nutrient (e.g., P only) reductions upstream may not necessarily reduce HAB impacts downstream. Reducing both N and P inputs is the only viable nutrient management solution for long-term control of HABs along the continuum. This article highlights where paired physical, chemical, or biological controls may improve beneficial uses in the short term, and offers management strategies that should be enacted across watershed scales to combat the global expansion of HABs across geographically broad freshwater-to-marine continua.

Coastal North Carolina (USA) has experienced 35 tropical cyclones over the past 2 decades; the frequency of these events is expected to continue in the foreseeable future. Individual storms had unique and, at times, significant hydrologic, nutrient-, and carbon (C)-loading impacts on biogeochemical cycling and phytoplankton responses in a large estuarine complex, the Pamlico Sound (PS) and Neuse River Estuary (NRE). Major storms caused up to a doubling of annual nitrogen and tripling of phosphorus loading compared to non-storm years; magnitudes of loading depended on storm tracks, forward speed, and precipitation in NRE-PS watersheds. With regard to C cycling, NRE-PS was a sink for atmospheric CO2 during dry, storm-free years and a significant source of CO2 in years with at least one storm, although responses were storm-specific. Hurricane Irene (2011) mobilized large amounts of previously-accumulated terrigenous C in the watershed, mainly as dissolved organic carbon, and extreme winds rapidly released CO2 to the atmosphere. Historic flooding after Hurricanes Joaquin (2015) and Matthew (2016) provided large inputs of C from the watershed, modifying the annual C balance of NRE-PS and leading to sustained CO2 efflux for months. Storm type affected biogeochemical responses as C-enriched floodwaters enhanced air–water CO2 exchange during ‘wet’ storms, while CO2 fluxes during ‘windy’ storms were largely supported by previously-accumulated C. Nutrient loading and flushing jointly influenced spatio-temporal patterns of phytoplankton biomass and composition. These findings suggest the importance of incorporating freshwater discharge and C dynamics in nutrient management strategies for coastal ecosystems likely to experience a stormier future.