Karl E. Havens

Controlling Eutrophication: Nitrogen and Phosphorus

Improvements in the water quality of many freshwater and most coastal marine ecosystems requires reductions in both nitrogen and phosphorus inputs.

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.

Rapid and highly variable warming of lake surface waters around the globe

In this first worldwide synthesis of in situ and satellite-derived lake data, we find that lake summer surface water temperatures rose rapidly (global mean = 0.34°C decade−1) between 1985 and 2009. Our analyses show that surface water warming rates are dependent on combinations of climate and local characteristics, rather than just lake location, leading to the counterintuitive result that regional consistency in lake warming is the exception, rather than the rule. The most rapidly warming lakes are widely geographically distributed, and their warming is associated with interactions among different climatic factors—from seasonally ice-covered lakes in areas where temperature and solar radiation are increasing while cloud cover is diminishing (0.72°C decade−1) to ice-free lakes experiencing increases in air temperature and solar radiation (0.53°C decade−1). The pervasive and rapid warming observed here signals the urgent need to incorporate climate impacts into vulnerability assessments and adaptation efforts for lakes.

Allied attack: climate change and eutrophication

Abstract Global warming and eutrophication in fresh and coastal waters may mutually reinforce the symptoms they express and thus the problems they cause.

Scale and Structure in Natural Food Webs

The degree to which widely accepted generalizations about food web structure apply to natural communities was determined through examination of 50 pelagic webs sampled consistently with even taxonomic resolution of all trophic levels. The fraction of species in various trophic categories showed no significant overall trends as the number of species varied from 10 to 74. In contrast, the number of links per species increased fourfold over the range of species number, suggesting that the link-species scaling law, defined on the basis of aggregated webs, does not reflect a real ecological trend.

ECOLOGY Controlling Eutrophication: Nitrogen and Phosphorus

Improvements in the water quality of many freshwater and most coastal marine ecosystems requires reductions in both nitrogen and phosphorus inputs.

Cyanobacteria blooms: effects on aquatic ecosystems

Cyanobacteria become increasingly dominant as concentrations of TP and TN increase during eutrophication of lakes, rivers and estuaries. Temporal dynamics of cyanobacteria blooms are variable--in some systems persistent blooms occur in summer to fall, whereas in other systems blooms are more sporadic. Cyanobacteria blooms have a wide range of possible biological impacts including potential toxic effects on other algae, invertebrates and fish, impacts to plants and benthic algae due to shading, and impacts to food web function as large inedible algae produce a bottleneck to C and energy flow in the plankton food web. In lakes with dense blooms of cyanobacteria, accumulation of organic material in lake sediments and increased bacterial activity also may lead to anoxic conditions that alter the structure of benthic macro-invertebrates. Diffusive internal P loading may increase, and hypolimnetic anoxia may lead to a loss of piscivorous fish that require a summer cold water refuge in temperate lakes. Ecosystem changes associated with frequent blooms may result in delayed response of lakes, rivers and estuaries to external nutrient load reduction. Despite numerous case studies and a vast literature on species-specific responses, community level effects of cyanobacterial blooms are not well understood--in particular the realized impacts of toxins and changes in food web structure/function. These areas require additional research given the prevalence of toxic blooms in the nations lakes, rivers and coastal waters--systems that provide a wide range of valued ecosystem services.

Complex interactions between autotrophs in shallow marine and freshwater ecosystems: implications for community responses to nutrient stress.

The relative biomass of autotrophs (vascular plants, macroalgae, microphytobenthos, phytoplankton) in shallow aquatic ecosystems is thought to be controlled by nutrient inputs and underwater irradiance. Widely accepted conceptual models indicate that this is the case both in marine and freshwater systems. In this paper we examine four case studies and test whether these models generally apply. We also identify other complex interactions among the autotrophs that may influence ecosystem response to cultural eutrophication. The marine case studies focus on macroalgae and its interactions with sediments and vascular plants. The freshwater case studies focus on interactions between phytoplankton, epiphyton, and benthic microalgae. In Waquoit Bay, MA (estuary), controlled experiments documented that blooms of macroalgae were responsible for the loss of eelgrass beds at nutrient-enriched locations. Macroalgae covered eelgrass and reduced irradiance to the extent that the plants could not maintain net growth. In Hog Island Bay, VA (estuary), a dense lawn of macroalgae covered the bottom sediments. There was reduced sediment-water nitrogen exchange when the algae were actively growing and high nitrogen release during algal senescence. In Lakes Brobo (West Africa) and Okeechobee (FL), there were dramatic seasonal changes in the biomass and phosphorus content of planktonic versus attached algae, and these changes were coupled with changes in water level and abiotic turbidity. Deeper water and/or greater turbidity favored dominance by phytoplankton. In Lake Brobo there also was evidence that phytoplankton growth was stimulated following a die-off of vascular plants. The case studies from Waquoit Bay and Lake Okeechobee support conceptual models of succession from vascular plants to benthic algae to phytoplankton along gradients of increasing nutrients and decreasing under-water irradiance. The case studies from Hog Island Bay and Lake Brobo illustrate additional effects (modified sediment-water nutrient fluxes, allelopathy or nutrient release during plant senescence) that could play a role in ecosystem response to nutrient stress.

Cyanobacterial toxins: a qualitative meta-analysis of concentrations, dosage and effects in freshwater, estuarine and marine biota.

This paper reviews the rapidly expanding literature on the ecological effects of cyanobacterial toxins. The study employs a qualitative meta-analysis from the literature examining results from a large number of independent studies and extracts general patterns from the literature or signals contradictions. The meta-analysis is set up by putting together two large tables--embodying a large and representative part of the literature (see Appendix A). The first table (Table A.1) reviews the presence (concentrations) of different cyanobacterial toxins in the tissues of various groups of aquatic biota after exposure via different routes, experimentally in the lab or via natural routes in the environment. The second table (Table A.2) reviews the dose dependent effect of toxins on biota. The great majority of studies deal with the presence and effects of microcystin, especially of the MC-LR congener. Although this may partly be justified--MC-LR is an abundant and highly toxic protein--our review also emphasizes what is known about (i) other MC congeners (a number of studies showed a preferred accumulation of the less toxic variant MC-RR in animal tissues), (ii) nodularin (data on a range of biota from studies on the Baltic Sea), (iii) neurotoxins like anatoxin-a(s), which are conspicuously often present at times when mass mortalities of birds occur, (iv) a few studies on the presence and effects of cylindrospermposin, as well as (v) the first examples of ecological effects of newly identified bioactive compounds, like microviridin-J. Data were reorganized to assess to what extent bioconcentration (uptake and concentration of toxins from the water) or biomagnification (uptake and concentration via the food) of cyanobacterial toxins occurs in ecosystems. There is little support for the occurrence of biomagnification, and this reduces the risk for biota at higher trophic levels. Rather than biomagnification biodilution seems to occur in the foodweb with toxins being subject to degradation and excretion at every level. Nevertheless toxins were present at all tropic levels, indicating that some vectorial transport must take place, and in sufficient quantities for effects to possibly occur. Feeding seemed to be the most important route for exposure of aquatic biota to cyanobacterial toxins. A fair number of studies focus on dissolved toxins, but in those studies purified toxin typically is used, and biota do not appear very sensitive to this form of exposure. More effects are found when crude cyanobacterial cell lysates are used, indicating that there may be synergistic effects between different bioactive compounds. Aquatic biota are by no means defenseless against toxic cyanobacteria. Several studies indicate that those species that are most frequently exposed to toxins in their natural environment are also the most tolerant. Protection includes behavioral mechanisms, detoxication of MC and NODLN by conjugation with glutathione, and fairly rapid depuration and excretion. A common theme in much of the ecological studies is that of modulating factors. Effects are seldom straightforward, but are dependent on factors like the (feeding) condition of the animals, environmental conditions and the history of exposure (acclimation and adaptation to toxic cyanobacteria). This makes it harder to generalize on what is known about ecological effects of cyanobacterial toxins. The paper concludes by summarizing the risks for birds, fish, macroinvertebrates and zooplankton. Although acute (lethal) effects are mentioned in the literature, mass mortalities of--especially--fish are more likely to be the result of multiple stress factors that co-occur during cyanobacterial blooms. Bivalves appear remarkably resistant, whilst the harmful effects of cyanobacteria on zooplankton vary widely and the specific contribution of toxins is hard to evaluate.

Zooplankton community responses to chemical stressors: A comparison of results from acidification and pesticide contamination research

The response of freshwater zooplankton communities to two chemical stressors, acidification and pesticide contamination, were investigated in a review of published research results. The objective was to test Odums predictions (Odum, 1985) that in response to stress, both the average body size of organisms and their efficiency in utilizing resources are reduced. Acidification and pesticide contamination were both found to favor dominance by small cladorecans and rotifers, the smallest zooplankton taxa. This finding was consistent with Odums predictions, however, there were exceptions to the trend. The dominance of small taxa may be due to rapid reproductive rates, physiological tolerance, development with few transitions through sensitive stages (eg. post-molting), or to the great richness of small species. Regardless of the mechanism, there is evidence that when acidification and pesticide contamination result in small zooplankton dominance, the efficiency of carbon and energy transfer from algae to zooplankton is reduced. This finding is also consistent with Odums predictions.