Bioaccumulation and trophic transfer of dioxins in marine copepods and fish

Despite the great concerns about dioxins in the marine environments, the biokinetics and bioaccumulation of these compounds in marine organisms remains little known. Using radioactive tracers the aqueous uptake, dietary assimilation efficiency, and elimination of dioxins were measured in marine phytoplankton, copepods and seabream. The calculated uptake rate constant of dioxins decreased with increasing trophic levels, whereas the dietary assimilation efficiency (AE) was 28.5-57.6% in the copepods and 36.6-70.2% in the fish. The dietary AE was highly dependent on the food concentrations and food type. The elimination rate constant of dioxin in the copepods varied with different exposure pathways as well as food concentration and food type. Biokinetic calculation showed that dietary accumulation was the predominant pathway for dioxin accumulation in marine copepods and fish. Aqueous uptake can be an important pathway only when the bioconcentration of dioxins in the phytoplankton was low.     

Effects of major nutrient additions on metal uptake in phytoplankton

We examined the influences of major nutrients (N, P, Si) on the accumulation of three trace metals [Cd, Se(IV), and Zn] in four species of marine phytoplankton (diatom, green alga, dinoflagellate, prasinophyte). Relative metal uptake was quantified by the kinetic measurements of metal concentration factor over a short exposure period. Our study demonstrated that nutrient addition significantly influenced the metal uptake rate and the cell growth rate in all four phytoplankton species. An increase in ambient N concentration considerably enhanced metal uptake by the cells. The dry weight concentration factor increased by 2.4-14.9 times for Cd, 1.1-4.0 times for Se, and 1.1-5.4 times for Zn in all four phytoplankton species with an addition of 176.4 microM N. The effects of P or Si addition on metal uptake and cell growth were less pronounced than the effects of N addition. Under most circumstances the rate of metal uptake increased exponentially with increasing cell growth rate constant. Only Se(IV) uptake in the diatom Phaeodactylum tricornutum was not correlated with cell growth rate. Se(IV) was not accumulated by the green algae Chlorella autotrophica at a high P concentration (7.2 microM), but appreciable accumulation was documented in cells inoculated without P addition. Our study therefore demonstrated that nutrient enrichments in many coastal waters can considerably affect trace metal uptake in phytoplankton and presumably metal trophic transfer in marine food chains.     

Uptake, absorption efficiency and elimination of DDT in marine phytoplankton, copepods and fish

Uptake, absorption efficiency and elimination of DDT were measured in marine phytoplankton, copepods (Acartia erythraea) and fish (mangrove snappers Lutjanus argentimaculatus). The uptake rate constant of DDT from water decreased with increasing trophic level. The dietary absorption efficiency (AE) of DDT was 10-29% in copepods and 72-99% in fish. Food concentration did not significantly affect the AEs of DDT, but the AEs varied considerably among the different food diets. The elimination rate constants of DDT by the copepods were comparable following uptake from the diet and from the water. Elimination of DDT from the fish was exceedingly low. Both aqueous and dietary uptake are equally important for DDT accumulation in the copepods. In fish, dissolved exposure is a more significant route than intake from the diet. The predicted trophic transfer factors in the copepods and the fish are consistent with the field measurements in marine zooplankton and fish.     

Biokinetics of cadmium, selenium, and zinc in freshwater alga Scenedesmus obliquus under different phosphorus and nitrogen conditions and metal transfer to Daphnia magna

The uptake of Cd, Se(IV) and Zn by the freshwater alga Scenedesmus obliquus and the subsequent transfer and release budget in Daphnia magna were investigated under different nutrient additions and cell incubation conditions. An increase in ambient phosphate concentrations from 0.5 micromol l(-1) to 50 micromol l(-1) significantly increased the intracellular accumulation of Cd (by 18x) and Zn (by 5x), but decreased the accumulation of Se (by 126x) in the alga. The percentage of these metals distributing in the intracellular pool of algae also increased substantially with increasing ambient P concentrations. Nitrate addition from 5.0 to 200 micromol l(-1) did not influence the uptake of any of the three metals, although a significant decrease in the intracellular Se distribution was observed. Radiolabeled algae under different nutrient manipulations (semi-continuous culture, starvation, and P-pulse treatments) were used to measure trophic transfer assimilation efficiency (AE) in Daphnia. When the algal cells were grown in a semi-continuous culture, starved for N and P, or were treated with P-pulse, the AEs of Cd and Zn were generally independent of the nutritional conditions, but the Se AE was significantly affected by different P levels. The efflux rate constants, determined during 10 d depuration following 7 days of dietary uptake, decreased significantly for Cd and Zn, but were relatively constant for Se with increasing P concentration. N-addition caused no effect on the metal efflux rate constants. P- or N-additions did not influence the release budget (including molting, neonates, excretion and feces) for all three elements in Daphnia. Our study indicated that phosphate enrichment may substantially increase metal uptake in green alga S. obliquus. Responses of trophic transfer in Daphnia to nutrient enrichment were metal specific. P-enrichment can possibly lead to considerable decrease on Se transfer from algae to zooplankton.     

Effects of macronutrient additions on nickel uptake and distribution in the dinoflagellate Prorocentrum donghaiense Lu

The influences of macronutrient additions on nickel (Ni) uptake and distribution in the subcellular structures and macromolecular components of the dinoflagellate Prorocentrum donghaiense Lu were examined using a radioisotope tracer method. The results showed that nitrate addition enhanced the uptake of Ni by P. donghaiense, whereas phosphate addition inhibited Ni uptake at high-Ni concentration. Nitrate or phosphate addition significantly affected Ni distribution in the subcellular structures and components. The majority of Ni was found in the soluble substances (>70%) and in the proteins (55.0-79.6%) of the algal cells. Urea reduced the Ni content in the amino acid-carbohydrate but elevated its content in proteins, and shown significantly correlated with the protein content of the algal cells. Thus, nutrient enrichment could influence both metal uptake and its distribution in the subcellular structures and components of the phytoplankton, as well as its subsequent transfer in marine food chains.     

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.     

Eutrophication of freshwater and coastal marine ecosystems a global problem

GOAL, SCOPE AND BACKGROUND: Humans now strongly influence almost every major aquatic ecosystem, and their activities have dramatically altered the fluxes of growth-limiting nutrients from the landscape to receiving waters. Unfortunately, these nutrient inputs have had profound negative effects upon the quality of surface waters worldwide. This review examines how eutrophication influences the biomass and species composition of algae in both freshwater and costal marine systems. MAIN FEATURES: An overview of recent advances in algae-related eutrophication research is presented. In freshwater systems, a summary is presented for lakes and reservoirs; streams and rivers; and wetlands. A brief summary is also presented for estuarine and coastal marine ecosystems. RESULTS: Eutrophication causes predictable increases in the biomass of algae in lakes and reservoirs; streams and rivers; wetlands; and coastal marine ecosystems. As in lakes, the response of suspended algae in large rivers to changes in nutrient loading may be hysteretic in some cases. The inhibitory effects of high concentrations of inorganic suspended solids on algal growth, which can be very evident in many reservoirs receiving high inputs of suspended soils, also potentially may occur in turbid rivers. Consistent and predictable eutrophication-caused increases in cyanobacterial dominance of phytoplankton have been reported worldwide for natural lakes, and similar trends are reported here both for phytoplankton in turbid reservoirs, and for suspended algae in a large river CONCLUSIONS: A remarkable unity is evident in the global response of algal biomass to nitrogen and phosphorus availability in lakes and reservoirs; wetlands; streams and rivers; and coastal marine waters. The species composition of algal communities inhabiting the water column appears to respond similarly to nutrient loading, whether in lakes, reservoirs, or rivers. As is true of freshwater ecosystems, the recent literature suggests that coastal marine ecosystems will respond positively to nutrient loading control efforts. RECOMMENDATIONS AND OUTLOOK: Our understanding of freshwater eutrophication and its effects on algal-related water quality is strong and is advancing rapidly. However, our understanding of the effects of eutrophication on estuarine and coastal marine ecosystems is much more limited, and this gap represents an important future research need. Although coastal systems can be hydrologically complex, the biomass of marine phytoplankton nonetheless appears to respond sensitively and predictably to changes in the external supplies of nitrogen and phosphorus. These responses suggest that efforts to manage nutrient inputs to the seas will result in significant improvements in coastal zone water quality. Additional new efforts should be made to develop models that quantitatively link ecosystem-level responses to nutrient loading in both freshwater and marine systems.     

Nitrogen in aquatic ecosystems

Aquatic ecosystems respond variably to nutrient enrichment and altered nutrient ratios, along a continuum from fresh water through estuarine, coastal, and marine systems. Although phosphorus is considered the limiting nutrient for phytoplankton production in freshwater systems, the effects of atmospheric nitrogen and its contribution to acidification of fresh waters can be detrimental. Within the estuarine to coastal continuum, multiple nutrient limitations occur among nitrogen, phosphorus, and silicon along the salinity gradient and by season, but nitrogen is generally considered the primary limiting nutrient for phytoplankton biomass accumulation. There are well-established, but nonlinear, positive relationships among nitrogen and phosphorus flux, phytoplankton primary production, and fisheries yield. There are thresholds, however, where the load of nutrients to estuarine, coastal and marine systems exceeds the capacity for assimilation of nutrient-enhanced production, and water-quality degradation occurs. Impacts can include noxious and toxic algal blooms, increased turbidity with a subsequent loss of submerged aquatic vegetation, oxygen deficiency, disruption of ecosystem functioning, loss of habitat, loss of biodiversity, shifts in food webs, and loss of harvestable fisheries.     

Photo-oxidation of Sb(III) in the seawater by marine phytoplankton-transition metals-light system

The photo-oxidation of Sb(III) to Sb(V) by marine microalgae (diatom, green and red algae) with or without the presence of transition metals (Fe(III), Cu(II) and Mn(II)). The influence of marine phytoplankton on the photochemistry of antimony was confirmed for the first time. The conversion ratio of Sb(III) to Sb(V) increased with increasing algae concentration and irradiation time. Different species of marine phytoplankton were found to have different photo-oxidizing abilities. The photochemical redox of transition metals could induce the species transformation of antimony. After photo-induced oxidation by marine phytoplankton and transition metals, the ratio of Sb(V) to Sb(III) was in the range of 1.07-5.48 for six algae (Tetraselmis levis, Chlorella autotrophica, Nannochloropsis sp., Tetraselmis subcordiformis, Phaeodactylum tricornutum, and Porphyridium purpureum), and only 0.92 for Dunaliella salina. The distribution of antimony in the sunlit surface seawater was greatly affected by combined effects of marine phytoplankton (main contributor) and transition metals; both synergistic and antagonistic effects were observed. The results provided further insights into the distribution of Sb(III) and Sb(V) and the biogeochemical cycle of antimony, and have significant implications for the risk assessment of antimony in the sunlit surface seawater.     

Nickel bioaccumulation in bivalves from the New Caledonia lagoon: seawater and food exposure

The New Caledonian lagoon is submitted to intense heavy metal input from land-based Ni mining. Therefore, the use of sentinel species is strongly recommended in order to develop and implement coastal zone management programmes of the metal contamination. The tropical oysters Isognomon isognomon and Malleus regula and the clam Gafrarium tumidum were previously proposed as such possible sentinel organisms and were thus investigated in this context. The three species were exposed to Ni via seawater or food using radiotracer techniques. Results indicate that uptake and retention efficiencies of Ni are independent of the dissolved Ni concentrations in the surrounding seawater. Hence, for the three species, body concentrations of Ni taken up from the dissolved phase are directly proportional to the ambient dissolved concentrations. Biokinetic patterns indicated that the major part of Ni was rapidly lost from bivalves during the first days of depuration, whereas 7 to 47% of 63Ni were retained in tissues with a biological half-life not significantly different from infinity. Finally, feeding experiments showed that Ni ingested with food (phytoplankton) was assimilated more efficiently in clams (assimilation efficiency, AE = 61%) than in oysters (AE = 17%), and was strongly retained (T(b1/2) > or = 35 d) in the tissues of both bivalves. It is concluded that the investigated species examined are efficient bioaccumulators of Ni from both the surrounding seawater and the food, and that they would be useful bioindicators for monitoring the status of Ni contamination in tropical coastal waters.     

Modelling the phytoplankton dynamics in a nutrient-rich solar saltern pond: predicting the impact of restoration and climate change

An ecological model for the solar saltern of Sfax (Tunisia) was established and validated by comparing simulation results to observed data relative to horizontal distributions of temperature, nutrients and phytoplankton biomass. Sensitivity analysis was performed in order to assess the influence of the main ecological model parameters. First applied at the saltern’s pond A1, the model was calibrated with field data measured over 4 years of study (from 2000 to 2003), which allowed an evaluation of parameters such as maximum growth rate of phytoplankton, optimal growth temperature and constant of half saturation for P/N assimilation by phytoplankton. Simulation results showed that the model allowed us to predict realistic phytoplankton variations of the study area, though we were unable to accurately reproduce the nutrient variation. The model was then applied to simulations of the impact of changes in phytoplankton biomass through scenarios such as hypothetic climate changes and saltern restoration.     

Accumulation and elimination of aqueous and dietary silver in Daphnia magna

The dissolved uptake, dietary assimilation, and efflux of Ag in a freshwater cladoceran, Daphnia magna, were measured under different laboratory conditions. The dissolved uptake rate of Ag was proportional to the ambient Ag concentration, but the accumulation was highly variable due to the sorption of Ag onto the daphnid bodies. The ambient Na(+) but not the ambient K(+) concentration significantly decreased the dissolved uptake of Ag, suggesting a competitive uptake of Ag(+) with Na(+). The dietary assimilation efficiencies (AEs) of Ag are dependent on the concentration of the algal food available to D. magna. The AE was as low as 2% when the food concentration reached the saturation levels. In contrast, the Ag concentrations in the algae did not significantly affect the Ag AE in D. magna. The efflux rate constant of diet-incorporated Ag was twice that through dissolved uptake. The elimination of Ag was further separated into different compartments (excretion, egestion, molting, and reproduction) in the juveniles and adults after accumulation from dissolved and dietary sources. Regeneration into the dissolved phase was the predominant pathway by which the incorporated Ag was lost from D. magna, regardless of the exposure pathway. In contrast to the essential metals or Hg, there was minimal maternal transfer of Ag from the mothers to the offspring. By employing the biokinetic model, we further showed that water is a dominant pathway for Ag accumulation in D. magna. Trophic transfer is less significant primarily because of the low Ag AE when the food concentration reached the saturation levels.     

Biotoxicity of nickel oxide nanoparticles and bio-remediation by microalgae Chlorella vulgaris

Adverse effects of manufactured nickel oxide nanoparticles on the microalgae Chlorellavulgaris were determined by algal growth-inhibition test and morphological observation via transmission electron microscopy (TEM). Results showed that the NiO nanoparticles had severe impacts on the algae, with 72 h EC(50) values of 32.28 mg NiOL(-1). Under the stress of NiO nanoparticles, C. vulgaris cells showed plasmolysis, cytomembrane breakage and thylakoids disorder. NiO nanoparticles aggregated and deposited in algal culture media. The presence of algal cells accelerated aggregation of nanoparticles. Moreover, about 0.14% ionic Ni was released when NiO NPs were added into seawater. The attachment of aggregates to algal cell surface and the presence of released ionic Ni were likely responsible for the toxic effects. Interestingly, some NiO nanoparticles were reduced to zero valence nickel as determined by X-ray diffraction (XRD) and X-ray photoelectron spectroscopy (XPS) analysis. The maximum ratios of nickel reduction was achieved at 72 h of exposure, in accordance with the time-course of changes in soluble protein content of treated C. vulgaris, implying that some proteins of algae are involved in the process. Our results indicate that the toxicity and bioavailability of NiO nanoparticles to marine algae are reduced by aggregation and reduction of NiO. Thus, marine algae have the potential for usage in nano-pollution bio-remediation in aquatic system.     

Nutrient retention in plant biomass and sediments from the salt marsh in Hangzhou Bay estuary, China

Nutrient load into the ocean can be retained during the process of plant uptake and sedimentation in marshes along the bay zone. Seasonal variations of biomass and nutrient concentration in three dominated plant assemblages and associated sediments were monitored in this study area to determine effects of salt marsh on nutrient retention. Results showed that plant aboveground biomass displayed a unimodal curve with nutrient concentration generally decreased from spring to winter. Belowground biomass was relatively low during the rapid growth period with nutrient concentration tending to decrease and then increase during this period. Plant total nitrogen (TN) pools are higher than total phosphorus (TP) pools, and both pools showed significant seasonal variations. Water purification coefficients (WPC) of nutrients by plant assimilation were 34.4/17.3, 19.3/24.0, and 5.14/6.04 t/(m² year) (TN/TP) for Phragmites australis, Spartina alterniflora, and Scirpus mariqueter, respectively. Overall, these results suggest that higher annual plant biomass and nutrient assimilation contribute to greater nutrient retention capacity and accumulation in sediments, thereby enabling reduced eutrophication in transitional waters.     

Biosorption of copper(II) from aqueous solutions by pre-treated biomass of marine algae Padina sp

Biosorption of heavy metals can be an effective process for the removal and recovery of heavy metal ions from aqueous solutions. The biomass of marine algae has been reported to have high uptake capacities for a number of heavy metal ions. In this paper, the adsorption properties of a pre-treated biomass of marine algae Padina sp. for copper(II) were investigated. Equilibrium isotherms and kinetics were obtained from batch adsorption experiments. The biosorption capacities were solution pH dependent and the maximum capacity obtained was 0.80 mmol/g at a solution pH of about 5. The biosorption kinetics was found to be fast, with 90% of adsorption within 15 min and equilibrium reached at 30 min. The effects of light metal ions on copper(II) uptake were studied and the presence of light metal ions did not affect copper(II) uptake significantly. Fixed-bed breakthrough curves for copper(II) removal were also obtained. This study demonstrated that the pre-treated biomass of Padina sp. could be used as an effective biosorbent for the treatment of copper(II) containing wastewater streams.     

The influence of EDDS on the uptake of heavy metals in hydroponically grown sunflowers

Phytoextraction is an environmentally friendly in situ technique for cleaning up metal contaminated land. Unfortunately, efficient metal uptake by remediation plants is often limited by low phytoavailability of the targeted metals. Chelant assisted phytoextraction has been proposed to improve the efficiency of phytoextraction. Phytoremediation involves several subsequent steps: transfer of metals from the bulk soil to the root surfaces, uptake into the roots and translocation to the shoots. Nutrient solution experiments address the latter two steps. In this context we investigated the influence of the biodegradable chelating agent SS-EDDS on uptake of essential (Cu and Zn) and non-essential (Pb) metals by sunflowers from nutrient solution. EDDS was detected in shoots and xylem sap for the first time, proving that it is taken up into the above ground biomass of plants. The essential metals Cu and Zn were decreased in shoots in the presence of EDDS whereas uptake of the non-essential Pb was enhanced. We suggest that in the presence of EDDS all three metals were taken up by the non-selective apoplastic pathway as the EDDS complexes, whereas in the absence of EDDS essential metal uptake was primarily selective along the symplastic pathway. This shows that synthetic chelating agents do not necessarily increase uptake of heavy metals, when soluble concentrations are equal in the presence and absence of chelates.     

Algae mediate submerged macrophyte response to nutrient and dissolved inorganic carbon loading: A mesocosm study on different species

Nutrient and dissolved inorganic carbon are two important factors that influence the development and species composition of submerged macrophyte communities in shallow lake ecosystems. Yet little is known about their potential interactive effects on the submerged macrophytes and competition outcome of macrophyte–phytoplankton. We performed a mesocosm experiment to investigate the growth and photosynthetic performance of three submerged macrophytes in relation to phytoplankton/epiphyton with nutrient and bicarbonate enrichment. During the experimental period (42 d), increase in nutrient loading in water column resulted in a substantial burst of phytoplankton and epiphyton growth and increased light attenuation. When combined with nutrient loading, bicarbonate enrichment also resulted in a heavily phytoplankton- and epiphyton-dominated state, although bicarbonate enrichment per se does not cause the phytoplankton and epiphyton growth. However, increase in nutrient loading in water column had significant negative impact on individual performances (growth and photosynthesis) of the three submerged macrophytes and bicarbonate enrichment increased the effect of eutrophication on two dissected-leaf macrophytes (M. spicatum and E. nuttallii). Furthermore, our results also suggest that species-specific photosynthetic performances occurred when submerged macrophytes in an environment with high abundance of phytoplankton/epiphyton. This study highlighted that increase in nutrient loading and bicarbonate in water column is likely to interactively impact both abiotic and biotic properties of a freshwater ecosystem. The interactions of these two factors could select macrophyte species that are able to resist the shading from phytoplankton/epiphyton, making these species more dominant in natural freshwater ecosystems.     

Application of canonical correspondence analysis to determine the ecological contribution of phytoplankton to PCBs bioaccumulation in Qinhuai River,Nanjing, China

The distribution and interactions of phytoplankton and 14 polychlorinated biphenyls (PCBs) were investigated using canonical correspondence analysis in autumn in the Qinhuai River, Nanjing, China. Concentrations of PCBs in water and algal samples ranged from 33.78 to 144.84 ng/L and from 0.21 to 19.66 ng/L (0.06 to 3.04 ng/mg biomass), respectively. The predominant residual species of PCBs in water samples were tri- through hexachlorobiphenyls, and the predominant residuals in algae were tri-, tetra-, and heptachlorobiphenyls. The degree of eutrophication affected phytoplankton composition and PCB bioaccumulation, and led to sample site- and algal species specificity of PCB residues in the study area. Chlorophyta, Bacillariophyta, and Euglenophyta had strong capacities to take up PCBs, whereas Cyanophyta was less involved in the transfer of these compounds. Bioaccumulation of PCBs by algae may be affected by water quality, chlorination, phytoplankton composition, and the structure of the PCBs and the algal cell walls.     

Trace metal accumulation by algae in Sepetiba Bay, Brazil

Trace metal concentrations (Zn, Cd, Pb, Cr and Cu) were determined in some benthic algae from Sepetiba Bay, Brazil. This region has been modified by the increase of industrial activities during the last 30 years. Among species sampled the brown algae Padina gymnospora contained the greatest amount of Zn. The Bioconcentration Factor for Zn in P. gymnospora was found to be 10(4) under laboratory conditions. The absorption of (65)Zn depended on exposure time and increased with Zn concentrations in the medium. An insignificant amount of Zn desorbed from algae suggested a very high affinity with cellular binding sites. Zinc uptake by P. gymnospora is discussed by considering field and laboratory conditions.     

Fate and effects of heavy metals in salt marsh sediments

The fate and effects of selected heavy metals were examined in sediment from a restored salt marsh. Sediment cores densely covered with Spartina patens were collected and kept either un-amended or artificially amended with nickel (Ni) under standardized greenhouse conditions. Ni-amendment had no significant effect on the fate of other metals in sediments, however, it increased root uptake of the metals. Metal translocation into the shoots was small for all metals. Higher Ni concentrations in plants from amended cores were accompanied by seasonal reductions in plant biomass, photosynthetic capacity and transfer efficiency of open photosystem II reaction centers; these effects, however, were no longer significant at the end of the growing season. Root colonization by arbuscular mycorrhizal fungi (AMF) resembled that of natural salt marshes with up to 20% root length colonized. Although Ni-amendment increased AMF colonization, especially during vegetative growth, in general AMF were largely unaffected.