Ammonium regeneration and assimilation in eelgrass (Zostera marina) beds

Regeneration and assimilation of ammonium in the water column and in sediments of eelgrass (Zostera marina L.) beds of Izembek Lagoon and Crane Cove, Alaska, USA and Mangoku-Ura, northeastern Japan, were investigated by using a ¹⁵N isotope dilution technique. In the water column of Mangoku-Ura, ammonium was regenerated at a rate of 12 nmol l^-1 h^-1 and assimilated at a rate of 74 nmol l^-1 h^-1. The ammonium regeneration rate in sediments ranged from 2 to 150 nmol g^-1 h^-1, and with one exception, exceeded ammonium assimilation in sediments (0.3 to 77 nmol g^-1 h^-1). The ammonium regeneration in the water column was of little significance for the nitrogen supply to the eelgrass bed ecosystem. Net ammonium production (regeneration minus assimilation) in the sediment of Izembek Laggon met nitrogen demand for eelgrass growth, suggesting that ammonium regeneration in the sediments was very important for the nitrogen cycle in the eelgrass bed ecosystem.     

Effects of anaerobiosis on root metabolism of Zostera marina (eelgrass): implications for survival in reducing sediments

The temperate seagrass Zostera marina L. typically grows in highly reducing sediments. Photosynthesis-mediated O₂ supplied to below-ground tissues sustains aerobic respiration during photosynthetic periods. Roots, however, experience daily periods of anoxia and/or hypoxia at night and under conditions that reduce photosynthesis. Rhizosphere cores of Z. marina were collected in August 1984 from Great Harbor, Massachusetts, USA. We examined short-term anaerobic metabolism of [U-¹⁴C]sucrose in excised roots and roots of intact plants. Under anaerobic conditions roots showed appreciable labeling of CO₂, ethanol and lactate, and slight labeling of alanine and other metabolites. Over 95% of the ¹⁴C-ethanol was recovered in the root exudate. Release of other metabolites from the roots was minimal. Ethanol was also released from hypoxic/anoxic roots of intact plants and none of this ethanol was transported to the shoot under any experimental conditions. Loss of ethanol from roots prevented tissue levels of this phytotoxin from increasing during anaerobiosis despite increased synthesis of ethanol. Anaerobic metabolism of [U-¹⁴C]glutamate in excised roots led to appreciable labelling of -aminobutyrate, which was known to accumulate in eelgrass roots. Roots recovered to fully aerobic metabolism within 4 h after re-establishment of aerobic conditions. The contributions of these root metabolic responses to the ability of Z. marina to grow in reducing marine sediments are related to light-regulated interactions of shoots and roots that likely dictate depth penetration, distribution and ecological success of eelgrass.     

Increased macrophyte nitrate reductase activity as a consequence of groundwater input of nitrate through sandy beaches

While most marine macrophytes preferentially assimilate ammonium to meet growth demand for nitrogen, some also utilize nitrate and exhibit high nitrate reductase activity (NRA). Although nitrate concentrations are often low in coastal waters during the summer and sandy beaches are generally considered to be low nutrient-input habitats, we have observed elevated NRA in leaves of some eelgrass (Zostera marina L.) plants growing immediately adjacent to the shoreline. We postulated that nitrate may become available to eelgrass and macroalgae via groundwater inputs that enter the nearshore water column. To address this possibility, we investigated the availability of groundwater nitrate for the induction of NRA in the leaves of eelgrass and in the macroalgaeSargassum filipendula C. Agardh (Phaeophyceae) andEnteromorpha intestinalis L. Link (Chlorophyceae) collected adjacent to two sandy beaches in the vicinity of Woods Hole, Massachusetts, USA. Induction of NRA was determined in the laboratory for eelgrass collected from one of the beach sites and from an offshore site, Lackey's Bay, which is isolated from groundwater input. At the two beach locations, pore water nitrate concentrations were 100 to 400µM within a few meters inland from the waterline. Nitrate efflux into the nearshore water column was quite high and variable (2160±660µmol m^–2 h^–1) when associated with rapid percolation (37±11 1 m^–2 h^–1) of nitrate-enriched pore water. Turbulent wave mixing rapidly diluted the nitrate. Macroalgae and eelgrass growing adjacent to a beach with high nitrate efflux had NR activities three- to sevenfold higher than those of algae and eelgrass growing along a beach section with low nitrate efflux. NRA of eelgrass plants from Lackey's Bay and Great Harbor increased in response to low daily nitrate additions (10 to 25µM) in the laboratory, with higher nitrate additions (50 to 200µM) yielding less dramatic responses. The increase in NRA was roughly three times higher for Great Harbor than for Lackey's Bay eelgrass. It appears that groundwater input of nitrate is sufficient to induce NRA in marine macrophytes growing near some beaches, including those with turbulent wave mixing.     

The nitrogen supply to intertidal eelgrass (Zostera marina)

The uptake of ammonium and nitrate by eelgrass (Zostera marina L.) was studied in two-compartment chambers. The plants were collected in 1992 from a population growing on a tidal flat in the S.W. Netherlands. They were incubated under conditions which reflected field conditions; this implied the use of natural seawater and sediment porewater as incubation media. In all six experiments, carried out over the course of a major part of the growing period (from July to the end of September), ammonium appeared to be much more important as a source of nitrogen than nitrate. The largest part was taken up by the leaves: uptake of ammonium by the leaves accounted for 68 to 92% of total plant nitrogen uptake. The uptake of nitrogen compounds by the root-rhizome system represented only 4 to 30% of total plant uptake. Thus, at least during flood tide, the leaves play the major role in nitrogen uptake in this intertidal population. During ebb tide, most of the plants are submerged in very shallow tidepools. It is suggested that during this phase of the tidal cycle, influx of porewater ammonium into the tide-pool water may enable the leaves to exploit local sediment resources.     

Nutrient regeneration in shallow-water sediments of the estuarine plume region of the nearshore Georgia Bight,USA

Benthic community respiration and the cycling of N and P were seasonally investigated in the unprotected, sandy sediments (Z5m) of the nearshore zone of the Georgia Bight, USA in 1981 and 1982. Nutrient exchange across the sediment-water interface was calculated from a diffusive model, measured by in-situ enclosure experiments and estimated from whole core incubations. Seasonally changing pore water profiles indicated that the sediments were not in steady-state with respect to N and P and showed the characteristics of enhanced interstitial water movement by benthic animals. Over an annual period the total flux of nitrogen measured in situ averaged 1812 mol m^-2 d^-1 from the sediments. NH ₄ ⁺ flux accounted for the vast majority of the total directly measured N flux (77%), followed by nitrate + nitrite (14%), and dissolved organic nitrogen (9%). Phosphorus flux averaged 537 mol m^-2 d^-1. A large ratio of in-situ fluxes to calculated diffusive fluxes (5.2:1) indicated flux enhancement due to benthic animal activity. ammonium fluxes measured in situ did not agree well with the rate of NH ₄ ⁺ produced in incubated whole cores (11.7 mmol m^-2 d^-1). Relative rates of C, N and P release throughout the year fluctuated considerably. Generally, nutrient fluxes were not simply related to respiration or temperature. As respiration was highly correlated with temperature, however, this suggested that respiration-regeneration was temporarily decoupled from exchange across the sediment-water interface. The annual C-N-P flux stoichiometry was 130:3.1:1. Using the rate at which NH ₄ ⁺ was produced in incubated cores the stoichiometry was 120:21:1. The anomalously low N flux measured in situ was attributed to a combination of denitrification and wave-and current-induced sediment nutrient flushing. The potential for sediment flushing is high as experiments showed that sediments were fluidized or resuspended down to 25 cm during large storms. Benthic nutrient flux contributed 40% to the annual P but only 11% to the annual N requirements of the pelagic primary producers.This is Contribution No. 558 from the University of Georgia Marine Institute. This work was supported by the Georgia Sea Grant College Program maintained by the National Oceanic and Atmospheric Administration, US Department of Commerce, under Grant No. NA80AA-D-00091     

Nitrate reductase activity in Zostera marina

Eelgrass (Zostera marina L.) has access to nutrient pools in both the water column and sediments. We investigated the potential for eelgrass to utilize nitrate nitrogen by measuring nitrate reductase (NR) activity with an in vivo tissue assay. Optimal incubation media contained 60 mM nitrate, 100 mM phosphate, and 0.5% 1-propanol at pH 7.0. Leaves had significantly higher NR activity than roots (350 vs 50 nmoles NO ₂ ^– produced g FW^–1 h^–1). The effects of growing depth (0.8 m MLW, 1.2 m, 3.0 m, 5.0 m) and location within the eelgrass meadow (patch edge vs middle) on NR activity were examined using plants collected from three locations in the Woods Hole area, Massachusetts, USA, in July 1987. Neither depth nor position within the meadow appear to affect NR activity. Nitrate enrichment experiments (200 M NO ₃ ^– for 6 d) were conducted in the laboratory to determine if NR activity could be induced. Certain plants from shallow depth (1.2 m) showed a significant response to enrichment, with NR activity increasing from >100 up to 950 nmoles NO ₂ ^– g FW^–1 h^–1 over 6 d. It appears that Z. marina growing in very shallow water (0.8 m) near a shoreline may be affected by ground water or surface run-off enrichments, since plants from this area exhibited rates up to 1 600 nmol NO ₂ ^– g FW^–1 h^–1. Water samples from this location consistently had slightly higher NO ₃ ^– concentrations (1.4 M) than all other collection sites (0.7 M). Thus, it is possible that chronic run-off or localized groundwater inputs can create sufficient NO ₃ ^– enrichment in the water column to induce nitrate reductase activity in Zostera leaves.     

The association of N2-fixing bacteria with sea urchins

Dinitrogen fixation associated with bacteria in the gastrointestinal tract of sea urchins appears to be a widespread phenomenon: sea urchins from the tropics (Diadema antillarum, Echinometra lacunter, Tripneustes ventricosus), the temperature zone (Strongylocentrotus droebachiensis) and the arctic (S. droebachiensis) exhibited nitrogenase activity (C₂H₂ reduction). Pronounced seasonal variation was found in nitrogenase activity of temperate sea urchins feeding on kelp (Laminaria spp.) and eelgrass (Zostera marina). The mean monthly nitrogenase activity was inversely correlated with the nitrogen content of the sea urchin's food, which varied up to fivefold over the course of a year. The highest rate of nitrogenase activity recorded for a temperate sea urchin during the 14 month sampling period was 11.6g N fixed g wet wt^-1 d^-1, with a yearly mean activity of 1.36 g N fixed g wet wt^-1 d^-1. Studies with ¹⁵N confirmed the C₂H₂ reduction results and showed incorporation of microbially-fixed nitrogen into S. droebachiensis demonstrating that N₂ fixation can be a source of N for the sea urchin. Laboratory experiments indicated that part of the sea urchin's (S. droebachiensis) normal gastrointestinal microflora is responsible for the observed nitrogenase activity.     

Eutrophication and trophic structure in response to the presence of the eelgrass <Emphasis Type="Italic">Zostera noltii</Emphasis>

In estuaries, eelgrass meadows contribute to fundamental ecosystem functions of estuaries, providing food to several predators and buffering the negative effects of eutrophication. We asked whether the presence of the eelgrass Zostera noltii decreased the nitrogen concentration in the overlying water, affected the sources of nitrogen sequestrated by primary producers and changed the benthic and pelagic food web structures. We also studied the importance of these food webs in providing food to fish. We compared bare sediment to sediment covered by a Z. noltii meadow, and examined nutrient concentrations in the water column and δ¹⁵N in primary producers as indicators of anthropogenic inputs of nutrients. We then measured both δ¹³C and δ¹⁵N in the tissues of plants and consumers to establish food web structures. There were no differences in the concentrations and sources of nitrogen between sites. Rather, δ¹⁵N values indicated anthropogenic inputs of N (e.g. sewage discharges, agriculture) in both sites. There were no major differences in the structure of the planktonic food web, which was in part sustained by particulate organic matter and supported most predator fish, and in the structure of the benthic food web. Nonetheless, there were differences in the sources of food for omnivore consumers and for the detritivore Scrobicularia plana. Overall, the benthic food web did not use food derived from the eelgrass or macroalgae deposited on the substratum. Suspension feeders used particulate and sediment organic matter, whereas the δ¹³C and δ¹⁵N values of the other consumers indicated a likely contribution of benthic microalgae. Furthermore, in both habitats we found large variability in the isotope signatures of benthic macrofauna consumers, which did not allow distinguishing clearly different trophic groups and indicated a high level of omnivory and a mixed diet opportunistically making use of the availability of food in the surroundings.     

Eelgrass fairy rings: sulfide as inhibiting agent

Distinct ‘fairy rings’ consisting of narrow fringes of eelgrass (Zostera marina L.) expand radially over a bottom of chalk plates outside the calcium carbonate cliffs of the island of Møn, Denmark. We conducted a survey to evaluate possible explanations for the formation of the rings and, more specifically, for the apparent die-off of eelgrass shoots on the inner side of the rings. The fairy rings were up to 15 m in diameter consisting of 0.3- to 1-m-wide zones of sea grass shoots at densities of up to 1,200 shoots m^?2 and rooted in an up to 10-cm-thick sediment layer. On the outer side, shoots expanded over the bare chalk plates. On the inner side, shoots were smaller, had lower absolute and specific leaf growth, shoot density was lower and the sediment eroded leaving the bare chalk with scattered boulders behind. Sediment organic matter and nutrients and tissue nutrient contents were not different among positions. Sediment pools of acid volatile sulfides and chromium-reducible sulfur increased from outer to the middle positions of the rings, and so did total sulfur content of eelgrass tissues, while tissue δ³⁴S isotope ratios, regardless of position in the fringes, were low reflecting substantial invasion of sulfide from the sediment. Neither the clonal growth pattern of eelgrass, sediment burial of shoots, hydrodynamic forcing nor nutrient limitation could explain the ring-shaped pattern. We conclude that the most likely explanation must be found in invasion of eelgrass shoots by toxic sulfide accumulating in the sediment due to low iron availability in the carbonate-rich environment.     

Reintroduction of eelgrass (Zostera marina) in the Dutch Wadden Sea; review of research and suggestions for management measures

Eelgrass (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal flats. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

Reintroduction of eelgrass ( Zostera marina ) in the Dutch Wadden Sea; Review of research and suggestions for management measures

Eelgras (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal falts. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

Genetic structure and gene flow of eelgrass <Emphasis Type="Italic">Zostera marina</Emphasis> populations in Tokyo Bay,Japan: implications for their restoration

Massive losses of eelgrass Zostera marina beds in Japan have occurred over the past 100 years. Toward their restoration, transplantation of eelgrass has been attempted in some areas, including Tokyo Bay. This study examined population genetic structures and gene flow in eelgrass in Tokyo Bay to establish guidelines for conducting restoration. Genotypes of a total of 360 individuals from 12 beds were determined using five microsatellite markers. The eelgrass beds in inner bay had above-average genetic diversity. A neighbor-joining tree based on F _ST values among beds revealed that a strong gene flow had occurred among six beds in the inner bay. Genetic assignment testing of drifting shoots indicated that those with seeds migrate in both directions between the inner and outer bay. We suggested that the restoration of eelgrass in the innermost part of Tokyo Bay, where natural habitats have been lost, should be conducted using the inner bay beds.     

Significance of salinity and silicon levels for growth of a formerly estuarine eelgrass (Zostera marina) population (Lake Grevelingen, The Netherlands)

Since the early 1980s, the eelgrass, Zostera marina L., population in the saline Lake Gevelingen, The Netherlands, is rapidly declining. An earlier study, in which long-term data on eelgrass coverage in this former estuary were correlated with several environmental variables, showed only one significant correlation: coverage was positively related to water column silicon levels. In addition, a negative correlation with salinity was observed, but this was not significant. In the present study, the effect of silicon and the effect of salinity on the development of Z. marina were investigated experimentally. Enhancement of dissolved silicon concentrations in the water did not stimulate Z. marina above-ground production or an increase in final above- and below-ground biomass. The highly significant correlation between eelgrass coverage and water column silicon levels, thus, remains to be explained. The results of the growth experiments did, however, demonstrate a clear effect of salinity on Z. marina growth. Plants cultured at 22 psu showed a higher production of shoots and leaves, resulting in more above-ground biomass, than plants grown at 32 psu. In addition, below-ground biomass was also higher at 22 psu. Measurements of chlorophyll a fluorescence, performed with a PAM-fluorometer, indicated a reduction of photosynthesis in the high-salinity treatments. Thus, low salinity stimulates development of Z. marina from Lake Grevelingen. Eelgrass from such a historically estuarine area may be more sensitive to high salinities than other, more marine populations. Recovery of the autochthonous eelgrass population is expected to be favoured when the estuarine conditions of the seagrass area are re-established, or when restoration programmes are carried out with allochthonous ecotypes that are less sensitive to high salinities. Received: 23 June 1998 / Accepted: 19 November 1998     

13C label identifies eelgrass (Zostera marina) Carbon in an Alaskan estuarine food web

The food web of Izembek Lagoon, Alaska draws most of its carbon from eelgrass (Zostera marina) and phytoplankton. The¹³C:¹²C ratios of these primary producers are sufficiently different to enable their contributions to consumers to be estimated from consumer¹³C:¹²C ratios. Although the technique is conceptually simple, carbon inputs from other sources and isotope fractionations occurring in the food web limit its precision. Isotopic data nevertheless helps to establish the major carbon fluxes through the community and to assess the importance of eelgrass carbon to individual animals. It is particularly useful when dealing with detritus food chains, where direct observations of animal feeding habits are difficult to make. The Izembek community draws much of its carbon from eelgrass. Detritus food chains provide the major pathway for assimilation of eelgrass carbon by the community, but grazers are also important. Eelgrass carbon is more important to benthic animals than to the eelgrass epibiota and the fishes, which depend mainly on phytoplankton carbon.Publication No. 381 of the Institute of Marine Science, University of Alaska.     

Sea level rise and eelgrass (Zostera marina) production: A spatially explicit relative elevation model for Padilla Bay, WA

The dynamics that govern the elevation of a coastal wetland relative to sea level are complex, involving non-linear feedbacks among opposing processes. Changes in the balance between these processes can result in significant alterations to vegetation communities that are adapted to a specific range of water levels. Given that current sedimentation rates in Padilla Bay, Washington are likely less than historical levels and that eustatic sea level rise is accelerating, the extensive Zostera marina (eelgrass) meadows in the bay may be at risk of eventual submergence. We developed a spatially explicit relative elevation model and used it to project changes in the productivity and distribution of eelgrass in Padilla Bay over the next century. The model is mechanistic and incorporates many of the processes and feedbacks that govern coastal wetland elevation change. Accretion estimates made using ²¹⁰Pb dating of sediment cores, sediment characteristics measured within cores, and eelgrass productivity and decomposition data were used to initialize and calibrate the model. Validation was performed using an elevation change rate measured with a network of surface elevation tables. Both the field data and model simulations revealed a net accretion deficit for the bay. Simulations using current rates of sea level rise indicated an overall expansion of eelgrass within Padilla Bay over the next century as it migrates from the center of the bay shoreward.     

Characterization of disjunct populations of Zostera marina (eelgrass) from California: genetic differences resolved by restriction-fragment length polymorphisms

Comparative restriction-fragment analysis was used to analyze the nuclear ribosomal DNA, and alcohol dehydrogenase-1 loci of Zostera marina L., for variation within and among populations. Eelgrass is a perennial marine flowering plant that is widespread and ecologically significant throughout the temperate northern hemisphere. A chemical method was developed to obtain restriction-quality DNA without CsCl fractionation from experimentally relevant quantities of seagrass tissues (0.5 to 1.0 g). The yield was 25 g g^-1 fresh weight. The three morphologically distinct forms of Z. marina from disjunct populations examined in this study were found to be genetically distinct; morphologically similar populations were indistinguishable genetically. Genetic distinction also correlated with habitat depth, as subtidal and intertidal populations were clearly divergent. Homologous probes for the 17S and 28S ribosomal DNA genes were used to map 24 restriction sites on the rDNA repeat of Z. marina, which was determined to be about 14 kb in length. At least 1 length mutation and 5 restriction-site changes were identified that distinguished Z. marina populations from San Diego and Monterey Bay (Del Monte Beach) from Z. marina populations from Elkhorn Slough and Tomales Bay. Estimated sequence variation (100×p) between eelgrass populations ranged from 0.00 to 0.69. Individual plants were observed to contain as many as four different rDNA-repeat length variants. The mean number of rDNA-repeat length variants per individual in Z. marina was about two. Intrapopulation variation in rDNA-repeat type was observed in only one individual from the Tomales Bay population.     

Production dynamics of the eelgrass, <Emphasis Type="Italic">Zostera marina</Emphasis> in two bay systems on the south coast of the Korean peninsula

Production dynamics of eelgrass, Zostera marina was examined in two bay systems (Koje Bay and Kosung Bay) on the south coast of the Korean peninsula, where few seagrass studies have been conducted. Dramatically reduced eelgrass biomass and growth have been observed during summer period on the coast of Korea, and we hypothesized that the summer growth reduction is due to increased water temperature and/or reduced light and nutrient availabilities. Shoot density, biomass, morphological characteristics, leaf productivities, and tissue nutrient content of eelgrass were measured monthly from June 2001 to April 2003. Water column and sediment nutrient concentrations were also measured monthly, and water temperature and underwater irradiance were monitored continuously at seagrass canopy level. Eelgrass shoot density, biomass, and leaf productivities exhibited clear seasonal variations, which were strongly correlated with water temperature. Optimal water temperature for eelgrass growth in the present study sites was about 15–20°C during spring period, and eelgrass growths were inhibited at the water temperature above 20°C during summer. Daily maximum underwater photon flux density in the study sites was usually much higher than the light saturation point of Z. marina previously reported. Densities of each terminal, lateral, and reproductive shoot showed their unique seasonal peak. Seasonal trends of shoot densities suggest that new eelgrass shoots were created through formation of lateral shoots during spring and a part of the vegetative shoots was transformed into flowering shoots from March. Senescent reproductive shoots were detached around June, and contributed to reductions of shoot density and biomass during summer period. Ambient nutrient level appeared to provide an adequate reserve of nutrient for eelgrass growth throughout the experimental period. The relationships between eelgrass growth and water temperature suggested that rapid reductions of eelgrass biomass and growth during summer period on the south coast of the Korean peninsula were caused by high temperature inhibition effects on eelgrass growth during this season.     

Seasonal variations in nitrogen-fixation (acetylene reduction) and sulphate-reduction rates in the rhizosphere of Zostera noltii: nitrogen fixation by sulphate-reducing bacteria

Nitrogen-fixation (acetylene reduction) rates were measured over an annual cycle in meadows of the seagrass Zostera noltii Hornem in the Bassin d'Arcachon, south-west France, between March 1994 and February 1995, using both slurry and whole-core techniques. Measured rates using the slurry technique consistently overestimated those determined on whole cores, probably due to the release of labile organic carbon sources as a result of root damage during preparation of the slurries. Thus, the whole-core technique may provide a more accurate estimate of in situ activity, since disturbance of physicochemical gradients of oxygen, sulphide, nutrients and the relationship between the plant roots and the rhizosphere microflora is minimised. Rates measured by the whole-core method were 1.8- to 4-fold greater (dependent upon season) in the light than those measured during dark incubations, indicating that organic carbon diffusing from the plant roots during photosynthesis was an important factor in regulating nitrogen fixation in the rhizosphere. Additions of sodium molybdate, a specific inhibitor of sulphate-reducing bacteria (SRB) inhibited acetylene-reduction activity by >80% as measured by both the slurry and whole-core techniques throughout the year, inferring that SRB were the dominant component of the nitrogen-fixing microflora. A mutualistic relationship between Z. noltii and nitrogen-fixing SRB in the rhizosphere, based on the exchange of organic carbon and fixed nitrogen is proposed. Acetylene- and sulphate-reduction rates showed distinct summer peaks which correlated with a reduced availability of ammonium in the sediment and the annual growth cycle of Z. noltii in the basin. Overall, these data indicate that acetylene reduction (nitrogen fixation) activity in the rhizosphere of Z. noltii was regulated both by the availability of organic carbon from the plant roots and maintenance of a low NH ₄ ⁺ concentration in the vicinity of the plant roots due to efficient assimilation of NH ₄ ⁺ by Z. noltii during the growth season. Nitrogenfixation rates determined from acetylene-reduction rates measured using the whole-core technique ranged from 0.1 to 7.3 mg N m^-2d^-1, depending on season, and were calculated to contribute between 0.4 and 1.1 g N m^-2yr^-1, or 6.3 to 12% of the annual fixed nitrogen requirement of Z. noltii.     

Control of microbial growth and of amphipod grazing by water-soluble compounds from leaves of Zostera marina

The role of minor components of the leaves of Zostera marina L. in altering the activity of micro-organisms directly (and indirectly by affecting amphipod grazers) was investigated in laboratory experiments, using plants collected at Roberts Bank (49°2N; 123°8W) on the west coast of Canada. Water-soluble extracts of eelgrass leaves inhibited the growth of a micro-alga (Platymonas sp.) and many marine bacteria at concentrations equivalent to as little as 10 mg dry leaf l^-1. The potency of leaf extracts was higher (1) in young, actively metabolizing tissue than in older leaves, and (2) in leaves collected during rapid growth in summer 1980 than during the following winter. Water-soluble inhibitors (especially phenolic acids) may explain the low biomass of epiphytes on actively growing leaves. Three phenolic acids inhibited the test micro-organisms at concentrations as low as 0.3 mg l^-1; caffeic acid was more potent than either protocatechuic or gentisic acid. Extracts of young leaves also inhibited grazing by amphipods [Eogammarus confervicolus (Stimpson)] on dead leaves. The time required for leaching of soluble inhibitors may account for the delay between the loss of leaves from the plants and the onset of grazing. Thus, interactions among the biotic components of the detritus ecosystem may be significantly modified by minor compounds in the leaves of Z. marina     

Ammonium excretion by gelationous zooplankton and their contribution to the ammonium requirements of microplankton in Chesapeake Bay

Ammonium excretion rates of recently collected specimens of gelatinous zooplankton, the scyphomedusan Chrysaora quinquecirrha DeSor and the etenophore Mnemiopsis leidyi A. Agassiz, were correlated with body mass and water temperature in measurements made from April to October 1989 and 1990. Rates ranged between 3.5 and 5.0 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for C. quinquecirrha and 3.0 to 4.9 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for M. leidyi. Excretion rate equations and in situ data on the size distributions and biomasses of gelatinous zooplankters and water temperature were used to estimate the contribution of ammonium by medusae and ctenophores to mesohaline Chesapeake Bay waters on several dates during April to October 1989 and 1990. We then compared the estimated contributions to direct measurements of ¹⁵NH ₄ ⁺ uptake by microplankton. The maximum estimated regeneration by gelatinous zooplankton was 5.8 g atoms NH ₄ ⁺ -N m^-3h^-1 at night in August 1990, when medusae biomass was greatest. This represents about 4% of the ammonium required by the microplankton. During the daytime on all dates, less than 1% of the ammonium required by microplanktion was supplied by gelatinous zooplankton. Therefore, gelatinous zooplankton appear to play a minor role in the ammonium cycle of Chesapeake Bay.