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2014, Science (New York, N.Y.)
AI
Algal blooms, driven by nutrient over-enrichment, have traditionally been managed by controlling phosphorus inputs. However, recent findings indicate that nitrogen plays a crucial role in algal proliferation, challenging the paradigm that phosphorus is the sole limiting factor. With increasing bioavailable nitrogen from fertilizers and waste, harmful cyanobacterial blooms often occur, impacting water quality and public health. Future research must address the contributions of nitrogen and redesign management strategies for freshwater ecosystems.
Hydrobiologia, 2019
Cyanobacterial harmful algal blooms (CyanoHABs) are an increasingly common feature of large, eutrophic lakes. Non-N 2-fixing CyanoHABs (e.g., Microcystis) appear to be proliferating relative to N 2-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., N 2 , NH 3 , N 2 O). Biological control on N inputs versus outputs, or the balance between N 2 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 colimitation and management strategy. The recent
Environmental Science & Technology, 2010
A worldwide proliferation of harmful (toxic, food-web altering, hypoxia-generating) algal blooms (HABs) has been linked to human nutrient (phosphorus (P) and nitrogen (N)) overenrichment. In addressing this pressing environmental and human health issue, reducing P inputs has received the most attention, in part because this nutrient was first identified as driving eutrophication (1), and because P is simpler and less expensive to remove from pollution sources (e.g., wastewater, industrial effluents). Furthermore, P reductions have been prescribed to control a particularly noxious group of HABs, the N 2 fixing cyanobacteria, which can obtain N 2 from the atmosphere, thereby minimizing ecosystem-level nitrogen (N) limitation
Harmful Algae
Western Lake Erie (WLE) experiences anthropogenic eutrophication and annual, toxic cyanobacterial blooms of non-nitrogen (N) fixing Microcystis. Numerous studies have shown that bloom biomass is correlated with an increased proportion of soluble reactive phosphorus loading from the Maumee River. Long term monitoring shows that the proportion of the annual Maumee River N load of non-nitrate N, or total Kjeldahl nitrogen (TKN), has also increased significantly (Spearman's ρ = 0.68, p = 0.001) over the last few decades and is also significantly correlated to cyanobacterial bloom biomass (Spearman's ρ = 0.64, p = 0.003). The ratio of chemically reduced N to oxidized N (TKN:NO3) concentrations was also compared to extracted chlorophyll and phycocyanin concentrations from all weekly sampling stations within WLE from 2009 to 2015. Both chlorophyll (Spearman's ρ = 0.657, p < 0.0001) and phycocyanin (Spearman's ρ = 0.714, p < 0.0001) were significantly correlated with TKN:NO3. This correlation between the increasing fraction of chemically reduced N from the Maumee River and increasing bloom biomass demonstrates the urgent need to control N loading, in addition to current P load reductions, to WLE and similar systems impacted by non-N-fixing, toxin-producing cyanobacteria.
SIL Proceedings, 1922-2010, 2000
Harmful Algae, 2016
Historically, phosphorus (P) has been considered the primary limiting nutrient for phytoplankton assemblages in freshwater ecosystems. This review, supported by new findings from Lake Erie, highlights recent molecular, laboratory, and field evidence that the growth and toxicity of some nondiazotrophic blooms of cyanobacteria can be controlled by nitrogen (N). Cyanobacteria such as Microcystis possess physiological adaptations that allow them to dominate low-P surface waters, and in temperate lakes, cyanobacterial densities can be controlled by N availability. Beyond total cyanobacterial biomass, N loading has been shown to selectively promote the abundance of Microcystis and Planktothrix strains capable of synthesizing microcystins over strains that do not possess this ability. Among strains of cyanobacteria capable of synthesizing the N-rich microcystins, cellular toxin quotas have been found to depend upon exogenous N supplies. Herein, multi-year observations from western Lake Erie are presented demonstrating that microcystin concentrations peak in parallel with inorganic N, but not orthophosphate, concentrations and are significantly lower (p < 0.01) during years of reduced inorganic nitrogen loading and concentrations. Collectively, this information underscores the importance of N as well as P in controlling toxic cyanobacteria blooms. Furthermore, it supports the premise that management actions to reduce P in the absence of concurrent restrictions on N loading may not effectively control the growth and/or toxicity of non-diazotrophic toxic cyanobacteria such as the cosmopolitan, toxin-producing genus, Microcystis.
Biogeosciences, 2018
Recent global water quality crises point to an urgent need for greater understanding of cyanobacterial harmful algal blooms (cHABs) and their drivers. Nearshore areas of Lake Erie such as Sandusky Bay may become seasonally limited by nitrogen (N) and are characterized by distinct cHAB compositions (i.e., Planktothrix over Microcystis). This study investigated phytoplankton N uptake pathways, determined drivers of N depletion, and characterized the N budget in Sandusky Bay. Nitrate (NO − 3) and ammonium (NH + 4) uptake, N fixation, and N removal processes were quantified by stable isotopic approaches. Dissimilatory N reduction was a relatively modest N sink, with denitrification, anammox, and N 2 O production accounting for 84, 14, and 2 % of sediment N removal, respectively. Phytoplankton assimilation was the dominant N uptake mechanism, and NO − 3 uptake rates were higher than NH + 4 uptake rates. Riverine N loading was sometimes insufficient to meet assimilatory and dissimilatory demands, but N fixation alleviated this deficit. N fixation made up 23.7-85.4 % of total phytoplankton N acquisition and indirectly supports Planktothrix blooms. However, N fixation rates were surprisingly uncorrelated with NO − 3 or NH + 4 concentrations. Owing to temporal separation in sources and sinks of N to Lake Erie, Sandusky Bay oscillates between a conduit and a filter of downstream N loading to Lake Erie, delivering extensively recycled forms of N during periods of low export. Drowned river mouths such as Sandusky Bay are mediators of downstream N loading, but climate-change-induced increases in precipitation and N loading will likely intensify N export from these systems.
Hydrobiologia, 2008
In the 1990s a sharp decrease in nitrogen loading occurred in Estonian rivers, bringing about a reduction of the nitrogen-to-phosphorus ratio (N:P ratio) in the large shallow lakes, Peipsi (3,555 km2, mean depth 7.1 m) and Võrtsjärv (270 km2, 2.8 m). The average mass ratio of total nitrogen (TN) and total phosphorus (TP) in Võrtsjärv (45) was about twice as high as that in Peipsi (22). In Peipsi, the N2-fixing Gloeotrichia echinulata, Aphanizomenon flos-aquae and Anabaena species prevailed in the summer phytoplankton, while in Võrtsjärv the dominant cyanobacteria were Limnothrix planktonica, L. redekei and Planktolyngbya limnetica, which cannot fix N2; the main N2-fixing taxa Aphanizomenon skujae and Anabaena sp. seldom gained dominance. In May–October the critical TN:TP mass ratio, below which N2-fixing cyanobacteria (Nfix) achieved high biomasses, was ∼40 in Võrtsjärv and ∼30 in Peipsi. The percentages of both total cyanobacteria (CY) and Nfix (CY% and Nfix%) in Peipsi achieved their maximum values at an N:P mass ratio at or below 20 for both TN:TP and Nmin:SRP. In Võrtsjärv, the TN:TP supporting a high Nfix% was between 30 and 40 and the Nmin:SRP supporting this high percentage was in the same range as that in Peipsi (not to implement nitrogen reduction measures without a simultaneous decrease of phosphorus concentration.
EDIS, 2022
Nitrogen and phosphorus are two nutrients that are essential for the growth and survival of plants and animals but are often present in short supply. Both nitrogen and phosphorus are applied regularly through fertilizer to increase the yield of crops needed to feed human populations and for residential and commercial landscaping purposes. This publication contains information for stakeholders, students, scientists, and environmental agencies interested in understanding how nitrogen and phosphorus affect water resources. Major revision by Ashley Smyth, H. Dail Laughinghouse IV, Karl Havens, and Thomas Frazer; 5 pp.https://edis.ifas.ufl.edu/sg118
Limnology and Oceanography, 2010
Lake Taihu (Taihu) is the third largest freshwater lake in China and an important drinking water, fishing, and tourism resource for Jiangsu Province. Recent toxic cyanobacterial blooms caused by excessive human nutrient loading have focused attention on arresting blooms and restoring the lake to acceptable water quality conditions by reducing nutrient inputs. Field sampling and in situ nutrient enrichment bioassays were conducted to determine seasonal patterns of nutrient limitation and nutrient thresholds for phytoplankton growth. The TN : TP and TDN : TDP mass ratios in the ambient water showed high seasonal variation and changed from 33-80 : 1 and 52-212 : 1, respectively, in winter and spring, and both declined to below 20 : 1 in summer. In spring and winter, total phytoplankton biomass and growth rates increased significantly with additions of P, with no primary effects from N, suggesting P limitation of phytoplankton growth. During the summer and fall bloom periods, however, N additions alone revealed a significant positive effect on phytoplankton growth, and P additions only stimulated phytoplankton growth once N had been added, suggesting that N was the primary limiting nutrient, with P being a secondarily limiting nutrient. When P enrichment was $ 0.20 mg P L 21 and N enrichment $ 0.80 mg N L 21 , growth of the toxin-producing, dominant bloom-forming cyanobacteria Microcystis spp. was not nutrient limited. This study suggests that availability of N during the summer is a key growth-limiting factor for the proliferation and maintenance of toxic Microcystis spp. blooms. Therefore, although P load reduction is important, N load reduction is essential for controlling the magnitude and duration of algal booms in Taihu.
PloS one, 2014
Excessive anthropogenic nitrogen (N) and phosphorus (P) inputs have caused an alarming increase in harmful cyanobacterial blooms, threatening sustainability of lakes and reservoirs worldwide. Hypertrophic Lake Taihu, China's third largest freshwater lake, typifies this predicament, with toxic blooms of the non-N2 fixing cyanobacteria Microcystis spp. dominating from spring through fall. Previous studies indicate N and P reductions are needed to reduce bloom magnitude and duration. However, N reductions may encourage replacement of non-N2 fixing with N2 fixing cyanobacteria. This potentially counterproductive scenario was evaluated using replicate, large (1000 L), in-lake mesocosms during summer bloom periods. N+P additions led to maximum phytoplankton production. Phosphorus enrichment, which promoted N limitation, resulted in increases in N2 fixing taxa (Anabaena spp.), but it did not lead to significant replacement of non-N2 fixing with N2 fixing cyanobacteria, and N2 fixation ...
Traditionally, nitrogen control is generally considered an important component of reducing lake eutrophication and cyanobacteria blooms. However, this viewpoint is refuted recently by researchers in China and North America. In the present paper, the traditional viewpoint of nitrogen control is pointed out to lack a scientific basis: the N/P hypothesis is just a subjective assumption; bottle bioassay experiments fail to simulate the natural process of nitrogen fixation. Our multi-year comparative research in more than 40 Yangtze lakes indicates that phosphorus is the key factor determining phytoplankton growth regardless of nitrogen concentrations and that total phytoplankton biomass is determined by total phosphorus and not by total nitrogen concentrations. These results imply that, in the field, nitrogen control will not decrease phytoplankton biomass. This finding is supported by a long-term whole-lake experiment from North America. These outcomes can be generalized in terms that a reduction in nitrogen loading may not decrease the biomass of total phytoplankton as it can stimulate blooms of nitrogen-fixing cyanobacteria. To mitigate eutrophication, it is not nitrogen but phosphorus that should be reduced, unless nitrogen concentrations are too high to induce direct toxic impacts on human beings or other organisms. Finally, details are provided on how to reduce controls on nitrogen and how to mitigate eutrophication.
Freshwater Biology, 2004
1. The sources of nitrogen for phytoplankton were determined for a bloom-prone lake as a means of assessing the hypothesis that cyanobacteria dominate in eutrophic lakes because of their ability to fix nitrogen when the nitrogen : phosphorous (N : P) supply ratio is low and nitrogen a limiting resource. 2. Nitrogen fixation rates, estimated through acetylene reduction with 15 N calibration, were compared with 15 N-tracer estimates of ammonium and nitrate uptake monthly during the ice-free season of 1999. In addition, the natural N stable isotope composition of phytoplankton, nitrate and ammonium were measured biweekly and the contribution of N 2 to the phytoplankton signature estimated with a mixing model. 3. Although cyanobacteria made up 81-98% of phytoplankton biomass during summer and autumn, both assays suggested minimal N acquisition through fixation (<9% for the in-situ incubations; <2% for stable isotope analysis). Phytoplankton acquired N primarily as ammonium (82-98%), and secondarily as nitrate (15-18% in spring and autumn, but <5% in summer). Heterocyst densities of <3 per 100 fixer cells confirmed low reliance on fixation. 4. The lake showed symptoms of both light and nitrogen limitation. Cyanobacteria may have dominated by monopolizing benthic sources of ammonium, or by forming surface scums that shaded other algae.
Carolina Digital Repository (University of North Carolina at Chapel Hill), 2016
Preventing harmful algal blooms (HABs) is needed to protect lakes and downstream ecosystems. Traditionally, reducing phosphorus (P) inputs was the prescribed solution for lakes, based on the assumption that P universally limits HAB formation. Reduction of P inputs has decreased HABs in many lakes, but was not successful in others. Thus, the "P-only" paradigm is overgeneralized. Whole-lake experiments indicate that HABs are often stimulated more by combined P and nitrogen (N) enrichment rather than N or P alone, indicating that the dynamics of both nutrients are important for HAB control. The changing paradigm from P-only to consideration of dual nutrient control is supported by studies indicating that (1) biological N fixation cannot always meet lake ecosystem N needs, and (2) that anthropogenic N and P loading has increased dramatically in recent decades. Sediment P accumulation supports long-term internal loading, while N may escape via denitrification, leading to perpetual N deficits. Hence, controlling both N and P inputs will help control HABs in some lakes and also reduce N export to downstream N-sensitive ecosystems. Managers should consider whether balanced control of N and P will most effectively reduce HABs along the freshwater-marine continuum.
Hydrobiologia, 2013
Phosphorus (P) is conventionally thought to limit production in freshwaters and nitrogen (N) that in the sea. Before much human activity, however, colimitation by N and P was probably normal, with systems developing ratios of N to P tending to the Redfield ratio. Single-factor limitation may be a symptom of human activity in many cases. It is widely believed that N fixation should compensate for N shortage because N fixers are ubiquitous and versatile, but this is not always the case and the issue has hitherto been considered largely with respect to plankton communities. Effects of N on macrophyte communities provide justification for control of both nutrients, at least in shallow lakes and estuaries. Increased N loading reduces plant biodiversity, changes the structure, and is associated with eventual loss, of macrophyte communities. P control alone may suffice in many deep lakes where denitrification is low and stratified conditions favour cyanobacterial development. Therein may lie a resolution to current controversies.
Proceedings of the National Academy of Sciences, 2008
Lake 227, a small lake in the Precambrian Shield at the Experimental Lakes Area (ELA), has been fertilized for 37 years with constant annual inputs of phosphorus and decreasing inputs of nitrogen to test the theory that controlling nitrogen inputs can control eutrophication. For the final 16 years (1990–2005), the lake was fertilized with phosphorus alone. Reducing nitrogen inputs increasingly favored nitrogen-fixing cyanobacteria as a response by the phytoplankton community to extreme seasonal nitrogen limitation. Nitrogen fixation was sufficient to allow biomass to continue to be produced in proportion to phosphorus, and the lake remained highly eutrophic, despite showing indications of extreme nitrogen limitation seasonally. To reduce eutrophication, the focus of management must be on decreasing inputs of phosphorus.
WIREs Water, 2019
Agricultural, urban and industrial activities have dramatically increased aquatic nitrogen and phosphorus pollution (eutrophication), threatening water quality and biotic integrity from headwater streams to coastal areas worldwide. Eutrophication creates multiple problems, including hypoxic "dead zones" that reduce fish and shellfish production; harmful algal blooms that create taste and odor problems and threaten the safety of drinking water and aquatic food supplies; stimulation of greenhouse gas releases; and degradation of cultural and social values of these waters. Conservative estimates of annual costs of eutrophication have indicated $1 billion losses for European coastal waters and $2.4 billion for lakes and streams in the United States. Scientists have debated whether phosphorus, nitrogen, or both need to be reduced to control eutrophication along the freshwater to marine continuum, but many management agencies worldwide are increasingly opting for dual control. The unidirectional flow of water and nutrients through streams, rivers, lakes, estuaries and ultimately coastal oceans adds additional complexity, as each of these ecosystems may be limited by different factors. Consequently, the reduction of just one nutrient upstream to control eutrophication can allow the export of other nutrients downstream where they may stimulate algal production. The technology exists for controlling eutrophication, but many challenges remain for understanding and managing this global environmental problem.
Environmental Science & Technology, 2011
2018
Recent global water quality crises point to an urgent need for greater understanding of cyanobacterial harmful algal blooms (cHABs) and their drivers. Nearshore areas of Lake Erie such as Sandusky Bay may become seasonally limited by nitrogen (N) and are characterized by distinct cHAB compositions (i.e., Planktothrix over Microcystis). This study investigated phytoplankton N uptake pathways, determined drivers of N depletion, and characterized the N budget in Sandusky Bay. Nitrate (NO − 3) and ammonium (NH + 4) uptake, N fixation, and N removal processes were quantified by stable isotopic approaches. Dissimilatory N reduction was a relatively modest N sink, with denitrification, anammox, and N 2 O production accounting for 84, 14, and 2 % of sediment N removal, respectively. Phytoplankton assimilation was the dominant N uptake mechanism, and NO − 3 uptake rates were higher than NH + 4 uptake rates. Riverine N loading was sometimes insufficient to meet assimilatory and dissimilatory demands, but N fixation alleviated this deficit. N fixation made up 23.7-85.4 % of total phytoplankton N acquisition and indirectly supports Planktothrix blooms. However, N fixation rates were surprisingly uncorrelated with NO − 3 or NH + 4 concentrations. Owing to temporal separation in sources and sinks of N to Lake Erie, Sandusky Bay oscillates between a conduit and a filter of downstream N loading to Lake Erie, delivering extensively recycled forms of N during periods of low export. Drowned river mouths such as Sandusky Bay are mediators of downstream N loading, but climate-change-induced increases in precipitation and N loading will likely intensify N export from these systems.
Water, 2013
The pronounced increase in the cycling and deposition of biologically reactive dissolved inorganic nitrogen (DIN) over large areas globally not only cause increased concentrations of DIN in surface waters, but it will also affect nutrient ratios in rivers, lakes and coastal areas. This review addresses the flux and fate of DIN, focusing NO 3 in lakes of boreal and alpine catchments. Not only DIN-deposition, but also catchment properties strongly affect the concentrations of NO 3 in lakes, as well as NO 3 :total P (TP) ratios. This ratio displays an extreme variability, and does also serve as an indicator of shift between N and P-limitation of aquatic autotrophs. A high share of forests and bogs in the catchment generally decreases NO 3 :total P ratios, while alpine and subalpine catchments with sparse vegetation cover may have high NO 3 :total P ratios, especially in regions with high DIN-deposition. Several empirical and experimental studies indicate a shift from an initial N to P-limitation, but for N-limited lakes, an increased growth of phytoplankton, periphytes and macrophytes may be accredited to elevated inputs of DIN. An intensified P-limitation may also be a consequence of elevated DIN-deposition. This P-limitation may again yield higher C:P-ratios in autotrophs with negative impacts on grazers and higher trophic levels.
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