Val H. Smith

NONPOINT POLLUTION OF SURFACE WATERS WITH PHOSPHORUS AND NITROGEN

Agriculture and urban activities are major sources of phosphorus and nitrogen to aquatic ecosystems. Atmospheric deposition further contributes as a source of N. These nonpoint inputs of nutrients are difficult to measure and regulate because they derive from activities dispersed over wide areas of land and are variable in time due to effects of weather. In aquatic ecosystems, these nutrients cause diverse problems such as toxic algal blooms, loss of oxygen, fish kills, loss of biodiversity (including species important for commerce and recreation), loss of aquatic plant beds and coral reefs, and other problems. Nutrient enrichment seriously degrades aquatic ecosystems and impairs the use of water for drinking, industry, agriculture, recreation, and other purposes. Based on our review of the scientific literature, we are certain that (1) eutrophication is a widespread problem in rivers, lakes, estuaries, and coastal oceans, caused by overenrichment with P and N; (2) nonpoint pollution, a major source of P and N to surface waters of the United States, results primarily from agriculture and urban activity, including industry; (3) inputs of P and N to agriculture in the form of fertilizers exceed outputs in produce in the United States and many other nations; (4) nutrient flows to aquatic ecosystems are directly related to animal stocking densities, and under high livestock densities, manure production exceeds the needs of crops to which the manure is applied; (5) excess fertilization and manure production cause a P surplus to accumulate in soil, some of which is transported to aquatic ecosystems; and (6) excess fertilization and manure production on agricultural lands create surplus N, which is mobile in many soils and often leaches to downstream aquatic ecosystems, and which can also volatilize to the atmosphere, redepositing elsewhere and eventually reaching aquatic ecosystems. If current practices continue, nonpoint pollution of surface waters is virtually certain to increase in the future. Such an outcome is not inevitable, however, because a number of technologies, land use practices, and conservation measures are capable of decreasing the flow of nonpoint P and N into surface waters. From our review of the available scientific information, we are confident that: (1) nonpoint pollution of surface waters with P and N could be reduced by reducing surplus nutrient flows in agricultural systems and processes, reducing agricultural and urban runoff by diverse methods, and reducing N emissions from fossil fuel burning; and (2) eutrophication can be reversed by decreasing input rates of P and N to aquatic ecosystems, but rates of recovery are highly variable among water bodies. Often, the eutrophic state is persistent, and recovery is slow.

Eutrophication: impacts of excess nutrient inputs on freshwater, marine, and terrestrial ecosystems

In the mid-1800s, the agricultural chemist Justus von Liebig demonstrated strong positive relationships between soil nutrient supplies and the growth yields of terrestrial plants, and it has since been found that freshwater and marine plants are equally responsive to nutrient inputs. Anthropogenic inputs of nutrients to the Earths surface and atmosphere have increased greatly during the past two centuries. This nutrient enrichment, or eutrophication, can lead to highly undesirable changes in ecosystem structure and function, however. In this paper we briefly review the process, the impacts, and the potential management of cultural eutrophication in freshwater, marine, and terrestrial ecosystems. We present two brief case studies (one freshwater and one marine) demonstrating that nutrient loading restriction is the essential cornerstone of aquatic eutrophication control. In addition, we present results of a preliminary statistical analysis that is consistent with the hypothesis that anthropogenic emissions of oxidized nitrogen could be influencing atmospheric levels of carbon dioxide via nitrogen stimulation of global primary production.

Eutrophication science: where do we go from here?

Cultural eutrophication has become the primary water quality issue for most of the freshwater and coastal marine ecosystems in the world. However, despite extensive research during the past four to five decades, many key questions in eutrophication science remain unanswered. Much is yet to be understood concerning the interactions that can occur between nutrients and ecosystem stability: whether they are stable or not, alternate states pose important complexities for the management of aquatic resources. Evidence is also mounting rapidly that nutrients strongly influence the fate and effects of other non-nutrient contaminants, including pathogens. In addition, it will be important to resolve ongoing debates about the optimal design of nutrient loading controls as a water quality management strategy for estuarine and coastal marine ecosystems.

N:P ratios, light limitation, and cyanobacterial dominance in a subtropical lake impacted by non-point source nutrient pollution

A long-term (28-year) data set was used to investigate historical changes in concentrations of phosphorus (P), nitrogen (N), N:P ratios, and Secchi disk transparency in a shallow subtropical lake (Lake Okeechobee, Florida, USA). The aim was to evaluate changes in the risk of N2-fixing cyanobacterial blooms, which have infrequently occurred in the lakes pelagic zone. Predictions regarding bloom risk were based on previously published N:P ratio models. Temporal trends in the biomass of cyanobacteria were evaluated using phytoplankton data collected in 1974, 1989-1992, and 1997-2000. Concentrations of pelagic total P increased from near 50 microg l-1 in the mid-1970s to over 100 microg l-1 in the late 1990s. Coincidentally, the total N:P (mass) ratio decreased from 30:1 to below 15:1, and soluble N:P ratio decreased from 15:1 to near 6:1, in the lake water. Published empirical models predict that current conditions favor cyanobacteria. The observations confirm this prediction: cyanobacteria presently account for 50-80% of total phytoplankton biovolume. The historical decrease in TN:TP ratio in the lake can be attributed to a decreased TN:TP ratio in the inflow water and to a decline in the lakes assimilation of P, relative to N. Coincident with these declines in total and soluble N:P ratios, Secchi disk transparency declined from 0.6 m to near 0.3 m, possibly due to increased mineral turbidity in the lake water. Empirical models predict that under the turbid, low irradiance conditions that prevail in this lake, non-heterocystous cyanobacteria should dominate the phytoplankton. Our observations confirmed this prediction: non-N2-fixing taxa (primarily Oscillatoria and Lyngbya spp.) typically dominated the cyanobacteria community during the last decade. The only exception was a year with very low water levels, when heterocystous N2-fixing Anabaena became dominant. In the near-shore regions of this shallow lake, low N:P ratios potentially favor blooms of N2-fixing cyanobacteria, but their occurrence in the pelagic zone is restricted by low irradiance and lack of stable stratification.

The ecology of algal biodiesel production

Sustainable energy production represents one of the most formidable problems of the 21st century, and plant-based biofuels offer significant promise. We summarize the potential advantages of using pond-grown microalgae as feedstocks relative to conventional terrestrial biofuel crop production. We show how pond-based algal biofuel production, which requires significantly less land area than agricultural crop-based biofuel systems, can offer additional ecological benefits by reducing anthropogenic pollutant releases to the environment and by requiring much lower water subsidies. We also demonstrate how key principles drawn from the science of ecology can be used to design efficient pond-based microalgal systems for the production of biodiesel fuels.

DEVELOPING NUTRIENT TARGETS TO CONTROL BENTHIC CHLOROPHYLL LEVELS IN STREAMS: A CASE STUDY OF THE CLARK FORK RIVER

Abstract Approaches for assessing the effects of lowering nutrients on periphyton biomass in streams and rivers are poorly developed in contrast to those for lakes. Here we present two complementary approaches to assess target nutrient concentrations in streams, given desired mean and maximum standing crops of benthic algal chlorophyll. In the first approach, a reference portion or reach of the river that typically exhibits acceptable levels of benthic chlorophyll is identified (i.e. seasonal mean and maximum values do not exceed desirable levels), and the target levels for instream nutrient concentrations are defined by mean nutrient levels in the reference region. In the second approach, regression and graphical analysis of a large stream database are used to identify acceptable levels of instream total N and total P. The first approach supplies site-specific nutrient targets, whereas the second places nutrient control into a broader, more comparative perspective. In order to link these target concentrations to specific nutrient control measures, we describe a spreadsheet model that can be used to translate changes in external loading by point sources into predicted new instream nutrient concentrations. These quantitative methods are applied here to the control of nuisance algal growth in the Clark Fork River, Montana. We suggest that, in general, maintenance of mean instream total N concentrations below 350 μg l−1 and total P below 30 μg l−1 will result in mean benthic algal chlorophyll a density below nuisance levels of 100 mg m−2 in most streams.

Chlorophyll-phosphorus relations in individual lakes. Their importance to lake restoration strategies.

The response of algal biomass to nutrient reduction is critically evaluated in 16 north temperate lakes by using data from the literature. The analysis confirms that reductions in total phosphorus concentration in the lakes are typically accompanied by consistent declines in chlorophyll. The data also suggest that this response can be expected whether a lake is phosphorus or nitrogen limited, although the magnitude of the response may differ. This is in contrast to the conclusion of a recent report which suggests that a threshold response is involved. Furthermore, the responses of some lakes appear unique and may not be accurately predicted by using current global eutrophication models. Modifications of these models to account for additional factors are urged, in order that these prediction errors may be decreased in the future.

CONSTRAINTS ON PRIMARY PRODUCER N:P STOICHIOMETRY ALONG N:P SUPPLY RATIO GRADIENTS

A current principle of ecological stoichiometry states that the nitrogen to phosphorus ratio (N:P) of primary producers should closely match that from environmental nutrient supplies. This hypothesis was tested using data from ponds in Michigan, USA, a freshwater mesocosm experiment, a synthesis of studies from diverse systems (cultures, lakes, streams, and marine and terrestrial environments), and simple dynamic models of producer growth and nutrient content. Unlike prior laboratory studies, the N:P stoichiometry of phytoplankton in Michigan ponds clustered around and below the Redfield ratio (7.2:1 by mass), despite wide variation in N:P supply ratios (2:1-63:1 by mass) and the presence of grazers. In a mesocosm experiment, the N:P stoichiometry of phytoplankton cells again deviated from a nearly 1:1 relationship with N:P supply. Phytoplankton seston exhibited lower N:P content than expected at high N:P supply ratios, and often higher N:P content than anticipated at low N:P supply ratios, regardless of herbivore presence. Similar devi- ations consistently occur in the N:P stoichiometry of algae and plants in the other diverse systems. The models predicted that both high loss rates (sinking, grazing) and physiological limits to nutrient storage capacity could attenuate producer stoichiometry. In the future, research should evaluate how limits to elemental plasticity of producers can influence the role of stoichiometry in structuring communities and ecosystem processes.

Industrial‐strength ecology: trade‐offs and opportunities in algal biofuel production

Microalgae represent one of the most promising groups of candidate organisms for replacing fossil fuels with contemporary primary production as a renewable source of energy. Algae can produce many times more biomass per unit area than terrestrial crop plants, easing the competing demands for land with food crops and native ecosystems. However, several aspects of algal biology present unique challenges to the industrial-scale aquaculture of photosynthetic microorganisms. These include high susceptibility to invading aquatic consumers and weeds, as well as prodigious requirements for nutrients that may compete with the fertiliser demands of other crops. Most research on algal biofuel technologies approaches these problems from a cellular or genetic perspective, attempting either to engineer or select algal strains with particular traits. However, inherent functional trade-offs may limit the capacity of genetic selection or synthetic biology to simultaneously optimise multiple functional traits for biofuel productivity and resilience. We argue that a community engineering approach that manages microalgal diversity, species composition and environmental conditions may lead to more robust and productive biofuel ecosystems. We review evidence for trade-offs, challenges and opportunities in algal biofuel cultivation with a goal of guiding research towards intensifying bioenergy production using established principles of community and ecosystem ecology.

RESPONSES OF A BACTERIAL PATHOGEN TO PHOSPHORUS LIMITATION OF ITS AQUATIC INVERTEBRATE HOST

Host nutrition is thought to affect the establishment, persistence, and severity of pathogenic infections. Nutrient-deficient foods possibly benefit pathogens by constraining host immune function or benefit hosts by limiting parasite growth and reproduction. However, the effects of poor elemental food quality on a hosts susceptibility to infection and disease have received little study. Here we show that the bacterial microparasite Pasteuria ramosa is affected by the elemental nutrition of its aquatic invertebrate host, Daphnia magna. We found that high food carbon:phosphorus (C:P) ratios significantly reduced infection rates of Pasteuria in Daphnia and led to lower within-host pathogen multiplication. In addition, greater virulent effects of bacterial infection on host reproduction were found in Daphnia-consuming P-deficient food. Poor Daphnia elemental nutrition thus reduced the growth and reproduction of its bacterial parasite, Pasteuria. The effects of poor host nutrition on the pathogen were further evidenced by Pasteurias greater inhibition of reproduction in P-limited Daphnia. Our results provide strong evidence that elemental food quality can significantly influence the incidence and intensity of infectious disease in invertebrate hosts.