Nitrogen fixation in the rhizosphere of marine angiosperms

High rates of acetylene reduction were observed in systems containing excised rhizomes of the Caribbean marine angiosperms Thalassia testudinum, Syringodium filiforme and Diplanthera wrightii, and the temperate marine angiosperm Zostera marina. For 4 plant and plant-sediment systems the ratio of acetylene reduced/N₂ fixed varied from 2.6 to 4.6. For T. testudinum the estimated rates of nitrogen fixation are in agreement with estimated requirements of the plant for nitrogen. For a typical T. testudinum stand, N₂ fixation is estimated to be 100 to 500 kg N/hectare per year. Numbers of N₂-fixing bacteria in the rhizosphere sediments were roughly 50 to 300 times more abundant than those in the nonrhizosphere sediments, and in both types of sediments were of the same orders as the estimated numbers of heterotrophic aerobes.Canadian IBP Contribution No. 137.     

Nitrogen fixation (acetylene reduction) associated with macroalgae in a coral-reef community in the Bahamas

N₂ fixation (C₂H₂ reduction) was associated with several species of macroalgae on a coral reef near Grand Bahama Island. The highest rates were associated with Microdictyon sp. (Chlorophyceae) and Dictyota sp. (Phaeophyceae). Extensive mats of filamentous blue-green algae, not heterotrophic bacteria, were the N₂ fixing agents: in experiments with samples of Microdictyon sp., the activity was lightdependent and not stimulated by organic compounds under either aerobic or anaerobic conditions. Assays in situ, at 20 m depth, and on shipboard, gave similar rates of N₂ fixation; the cyanophytes presumably have pigment adaptations to function in blue light. The maximum rate of N₂ fixation, associated with Microdictyon sp., was 3.8 g N fixed g dry weight^-1 h^-1. Coral-reef communities flourish in nutrientimpoverished waters, and therefore any input of nitrogen is probably important in stabilizing such ecosystems.     

Nitrogen fixation (acetylene reduction) associated with the living coral Acropora variabilis

To estimate N₂-fixation, acetylene reduction assays were carried out on portions of the branches of the coral Acropora variabilis from the west coast of Malaysia. In some experiments, a sub-surface incubation apparatus was employed that was designed to keep the coral fragments near to their natural depth of occurrence. Other shipboard experiments used metabolic inhibitors to investigate the class of organism reducing acetylene. Stumps of coral gave the highest rates of activity, probably attributable to loosely associated cyanophytes. Coral tips also reduced acetylene at relatively high rates; reduction was enhanced in light by increased CO₂ concentration and decreased O₂ tensions indicative of photosynthetic bacteria. Algal material was not obvious on the tip surfaces and so the active organism was probably more integral to the coral structure than it was in the stumps. Maximum rates of acetylene reduction measured translated to 2.5 mg N₂ fixed per outcrop per day.     

Effects of enhanced UVB radiation on a marine benthic diatom mat

In order to study the ecophysiological response of a natural diatom-dominated microbial mat to an enhanced level of ultraviolet-B radiation (UVBR), intact sediment cores from a shallow microtidal bay on the Swedish west coast were incubated in an outdoor flow-through system and exposed to either no UVBR or to an artificially enhanced level of UVBR. The microbial mat was cohesive and dominated by the large (≈ 400 μm) motile diatom Gyrosigma balticum. Functional response to UVBR was assessed by measuring carbon fixation (¹⁴C) and allocation of photosynthetic products. Sediment oxygen microprofiles were measured as indicators of the balance between the photosynthetic and respiratory activity of the community. Structural variables included biomass and composition of microalgae, pigment composition (HPLC), content of UV-absorbing compounds, and composition of fatty acids. An enhanced level of UVBR resulted in significant functional changes in the microalgal community. Significantly decreased carbon fixation and lower net oxygen production (as calculated from oxygen microprofiles) suggest that primary productivity in the type of microbial mat studied is potentially sensitive to an enhanced level of UVBR. The pattern of carbon allocation hinted a change as to the photosynthetically active part of the community when exposed to UVBR, indicating a differential sensitivity among algal cells. Despite significant functional effects, pigment or algal composition were not significantly affected by enhanced UVBR. The concentration of UV-absorbing compounds was low and did not increase at UVBR exposure. Due to the short duration of the experiment (4 d), a change in algal biomass or composition was not to be expected. Moreover, the thickness of the sampled sediment layer, may have contributed to the lack of observed structural effects of enhanced UVBR. Visual observations suggested that UVBR affected the motility of G. balticum. This study, as well as other experiments with a similar diatom mat, suggest that vertical migration is a key mechanism to be further studied in relation to UVBR exposure of diatom-dominated microbial mats. Received: 7 September 1996 / Accepted: 8 October 1996     

Regime shift in a coastal marine ecosystem

We demonstrate changes in ecosystem stable states in a coastal lagoon that are consistent with what a regime shift would hypothesize. In the nutrient-stressed Ringk?bing Fjord, Denmark, a small change in one variable (salinity) facilitated by a change in sluice management, caused a sudden regime shift from a bottom-up controlled turbid state, into a top-down controlled clear-water state. The change in dominating pathway of organic matter production from pelagic turnover to benthic-pelagic coupling was facilitated by new recruitment and growth of existing suspension-feeding clams, Mya arenaria. With the invasion of clams, benthic grazing became the key feature of the biological structure. Phytoplankton composition and zooplankton abundance were also affected by the change in biological structure. The small, but sudden, increase in salinity caused by the change in sluice management led to a dramatic reduction in biomass and coverage of benthic vegetation and thus affected herbivorous waterbird populations. In recent years, plant coverage has been increasing, as can be expected with increased water transparency. The regime shift has some major implications for coastal water management and revealed some conflicts between different types of nature and environmental protection legislation.     

Nitrogen fixation on a coral reef

Acetylene reduction was used to assess nitrogen fixation on all major substrates at all major areas over a period of 1 to 6 yr (1980–1986) at One Tree Reef (southern Great Barrier Reef). Experiments using ¹⁵N₂ gave a ratio of 3.45:1.0 for C₂H₂ reduced:N₂ fixed. Acetylene reduction was largely light-dependent, saturated at 0.15 ml C₂H₂ per ml seawater, and linear over 6 h. High fixation was associated with two emergent cyanophyte associations, Calothrix crustacea and Scytonema hofmannii, of limited distribution. Subtidally, the major contribution to nitrogen fixation came from well-grazed limestone substrates with an epilithic algal community in the reef flat and patch reefs (3 to 15 nmol C₂H₄ cm^-2 h^-1). Similar substrates from the outer reef slope showed lower rates. Nitrogen fixation on beach rock, intertidal coral rubble, reef crest and lagoon sand was relatively small (0.3 to 1.0 nmol C₂H₄ cm^-2 h^-1). Seasonal changes in light-saturated rates were small, with slight reduction only in winter. Rates are also reported for experimental coral blocks (13 to 39 nmol cm^-2 h^-1) and for branching coral inside and outside territories of gardening damselfish (3 to 28 nmol cm^-2 h^-1). This work supports the hypothesis that the high nitrogen fixation on the reef flat and patch reefs of the lagoon (34 to 68 kg N ha^-1 yr^-1) is because these subtidal areas support highly disturbed communities with the greatest abundance of nitrogen-fixing cyanophyte algae. It is calculated from a budget of all areas that One Tree Reef has an annual nitrogen fixation rate of 8 to 16 kg N ha^-1 yr^-1.     

Nitrogen cycling in microbial mats: rates and patterns of denitrification and nitrogen fixation

Spatial and temporal variations in nitrogen fixation and denitrification rates were examined between July 1991 and September 1992 in the intertidal regions of Tomales Bay (California, USA). Microbial mat communities inhabited exposed mudflat and vegetated marsh surface sediments. Mudflat and marsh sediments exhibited comparable rates of nitrogen fixation. Denitrification rates were higher in marsh sediments. Nitrogen fixation rates were lowest during January at both sites, whereas highest rates occurred during summer and fall. Denitrification rates were highest during fall and winter months in marsh sediments, while rates in mudflat sediments were highest during summer and fall. In mudflat sediments, nitrogen fixation and denitrification rates, integrated over 24 h, ranged from 6 to 79 mg N m^-1 d^-1 and 1 to 10 mg N m^-2 d^-1, respectively. Rates of denitrification represented between 6 and 20% of nitrogen fixation rates during the day, but exceeded or were equivalent to nitrogen fixation rates at night. The highest integrated rates of both nitrogen fixation and denitrification occurred during July, whereas, the highest percent loss occurred during spring when denitrification rates amounted to 20% of nitrogen fixation rates during the day. Over an annual cycle, inputs of fixed N to mudflat communities occurred exclusively during daylight. These results underscore the importance of determining integrated diel rates of both nitrogen fixation and denitrification when constructing N budgets. Using this approach, it was shown that microbial denitrification can represent a significant loss of combined nitrogen from mats on daily as well as monthly time scales.     

Nitrogen fixation by blue-green algae associated with the siphonous green seaweed Codium decorticatum: effects on ammonium uptake

Nitrogen fixation (acetylene reduction) at rates of up to 1.2 g N₂ g dry wt^-1 h^-1 was measured for the siphonous green seaweed Codium decorticatum. No nitrogenase activity was detected in C. isthmocladum. The nitrogenase activity was light sensitive and was inhibited by the addition of DCMU and triphenyl tetrazolium chloride. Additions of glucose did not stimulate nitrogen fixation. Blue-green algae (Calothrix sp., Anabaena sp., and Phormidium sp.) were implicated as the organisms responsible for the nitrogenase activity. They occurred in a reduced microzone within the C. decorticatum thallus where nitrogen fixation was optimized. Nitrogen fixation did not affect the kinetic constants for ammonium uptake in C. decorticatum (K_s=12.0 M, V_max=13.4 mol NH₃ g dry wt^-1 h^-1) determined using the perturbation method. Nevertheless, C. decorticatum thalli which fixed nitrogen had internal dissolved nitrogen concentrations which were over 1.4 times higher than in non-fixing thalli. This suggests that if C. decorticatum does derive part of its nitrogen requirement from the blue-green algae which it harbors, the transfer does not involve competition between this process and the uptake of ambient ammonium.     

Thraustochytrid protists as a component of marine microbial films

Although ubiquitous in the marine environment, the presence and importance of thraustochytrid fungoid protists in primary film formation of freshly immersed surfaces in the sea have not been investigated. We isolated thraustochytrids from surfaces of glass, aluminium, mild steel and fibre glass panels immersed in sea water for 1 to 4 d, but not from those of copper and cupro-nickel. Microscopic examination of the primary film on glass surfaces revealed the presence of thraustochytrids. Thraustochytrids grew to varying population densities on surfaces of glass, aluminium and fibre glass in the laboratory. Scanning electron microscopy of the growth of Ulkenia profunda Gaertner, isolated from the primary film, on surfaces of glass and aluminium suggested that cells of the thraustochytrid attached directly to the surfaces, without producing copious extracellular polysaccharides. The presence of ectoplasmic net elements was not a prerequisite for attachment of cells to surfaces. Cell surface hydrophobicity of the thraustochytrid, as estimated by the MATH assay, might play an important role in adhesion. The presence of thraustochytrid cells on a polystyrene surface markedly induced settlement of barnacle larvae (Balanus amphitrite), as compared to barnacle extract and a control. Received: 26 July 1999 / Accepted: 21 January 2000     

Density-dependent migratory waves in the marine pelagic ecosystem

The migration of large aggregations of animals that sweep through the landscape is a phenomenon with large consequences in many ecosystems. It has been suggested that such migrations are mediated by resource depletion. Under this hypothesis it has been shown that simple foraging rules may generate density-dependent migratory waves (DDMW) in which the speed and amplitude increase with animal abundance. We tested these predictions on a 32-year data set of the spatial distribution of the two youngest age groups of a small pelagic schooling fish, the capelin (Mallotus villosus), by the end of their annual feeding migration in the Barents Sea. Our data suggest that the two age groups divided the Barents Sea by forming migratory waves that moved in opposite directions. The aggregation and spatial displacement of these waves increased with increasing age-specific abundance. However, possibly through social interactions, migratory pattern was modified by the abundance of the other age group.     

Nitrogen dynamics in the sediments of a wetland coastal ecosystem of southern India

The study area, Kuttanad Waters is a part of the Cochin estuarine system on the west coast of India. Kuttanad is well known for its agricultural activity and so the major contribution to the inorganic ions of nitrogen will be from fertilisers applied in agriculture. Based on observed salinity the stations have been divided into three zones. The fresh water zones had higher quantities of silt and clay whereas the estuarine zone was more sandy. The chemical speciation scheme applied here distinguishes three forms of ammoniacal nitrogen species: exchangeable, fixed, and organic ammoniacal nitrogen. No significant trends were observed in the seasonal distribution of total, exchangeable, fixed and organic nitrogen. A significant concentration of exchangeable ammonia was observed in the sediment due to their predominantly reducing environment, which restricts nitrification. High NH₄-N concentrations in the pore waters, along with the sedimentary composition leads to a significantly high quantity of fixed NH₄-N. The low values for N org is due to high mineralisation or deamination of organic nitrogen     

Nitrogen dynamics in the sediments of a wetland coastal ecosystem of southern India

The study area, Kuttanad Waters is a part of the Cochin estuarine system on the west coast of India. Kuttanad is well known for its agricultural activity and so the major contribution to the inorganic ions of nitrogen will be from fertilisers applied in agriculture. Based on observed salinity the stations have been divided into three zones. The fresh water zones had higher quantities of silt and clay whereas the estuarine zone was more sandy. The chemical speciation scheme applied here distinguishes three forms of ammoniacal nitrogen species: exchangeable, fixed, and organic ammoniacal nitrogen. No significant trends were observed in the seasonal distribution of total, exchangeable, fixed and organic nitrogen. A significant concentration of exchangeable ammonia was observed in the sediment due to their predominantly reducing environment, which restricts nitrification. High NH₄-N concentrations in the pore waters, along with the sedimentary composition leads to a significantly high quantity of fixed NH₄-N. The low values for N org is due to high mineralisation or deamination of organic nitrogen     

Meta-analysis of ecosystem services associated with oyster restoration

Restoration of foundation species promises to reverse environmental degradation and return lost ecosystem services, but a lack of standardized evaluation across projects limits understanding of recovery, especially in marine systems. Oyster reefs are restored to reverse massive global declines and reclaim valuable ecosystem services, but the success of these projects has not been systematically and comprehensively quantified. We synthesized data on ecosystem services associated with oyster restoration from 245 pairs of restored and degraded reefs and 136 pairs of restored and reference reefs across 3500 km of U.S. Gulf of Mexico and Atlantic coastlines. On average, restoration was associated with a 21-fold increase in oyster production (mean log response ratio = 3.08 [95% confidence interval: 2.58–3.58]), 34–97% enhancement of habitat provisioning (mean community abundance = 0.51 [0.41–0.61], mean richness = 0.29 [0.19–0.39], and mean biomass = 0.69 [0.39–0.99]), 54% more nitrogen removal (mean = 0.43 [0.13–0.73]), and 89–95% greater sediment nutrients (mean = 0.67 [0.27–1.07]) and organic matter (mean = 0.64 [0.44–0.84]) relative to degraded habitats. Moreover, restored reefs matched reference reefs for these ecosystem services. Our results support the continued and expanded use of oyster restoration to enhance ecosystem services of degraded coastal systems and match many functions provided by reference reefs.     

Nitrogen fixation in the North Pacific Ocean

Nitrogen fixation in the euphotic zone of the ocean was measured by C₂H₂ reduction and ¹⁵N₂ incorporation associated with Trichodesmium sp. and also with Richelia intracellularis occurring within the cells of Rhizosolenia styliformis var. longispina, and R. cylindrus. The vertical distribution of N₂ fixation activity, N₂-fixing species, particulate matter and dissolved nutrients was measured. The effects of light intensity, sample concentration, length of incubation, and nutrient enrichment on the rates of C₂H₂ reduction were determined. Estimates of the importance of N₂ fixation in adding previously uncycled nitrogen to the euphotic zone are given.     

Nitrogen exchange between a southeastern USA salt marsh ecosystem and the coastal ocean

The salt marsh ecosystem at North Inlet, South Carolina, USA consistently exported dissolved inorganic nitrogen via tidal exchange with the coastal Atlantic Ocean. Concentrations centrations of NH ₄ ⁺ and NO ₃ ^- +NO ₂ ^- displayed distinct tidal patterns with rising values during ebb flow. These patterns suggest the importance of biogeochemical processes in the flux of material from the salt marsh. NH ₄ ⁺ export peaked during the summer (15 to 20 mg m^-2 tide^-1) during a net balance of tidal water exchange. Remineralization of NH ₄ ⁺ within the salt marsh system appears to be contributing to the estimated annual net export of bout 4.7 g NH ₄ ⁺ -N m^-2 yr^-1. NO ₃ ^- +NO ₂ ^- exports were higher in the fall and winter of 1979 (2 to 4 mg N m^-2 tide^-1). The winter export coincided with a considerable net export of water with no distinctive concentration patterns, suggesting a simple advective export. However, the fall peak of NO ₃ ^- +NO ₂ ^- export occurred during a period of net water balance in tidal exchange and an insignificant freshwater input from the western, forested boundary. During the summer and fall, tidal concentration patterns were particularly apparent, suggesting that nitrification within the salt marsh system was contributing to the estimated annual net export of ca 0.6 g NO ₃ ^- +NO ₂ ^- -N m^-2 yr^-1.Contribution No. 637 from the Belle W. Baruch Institute of Marine Biology and Coastal Research     

Nitrogen mineralization in a high altitude ecosystem in the mediterranean phytogeographical region of Turkey

Interrelations exist in the terrestrial ecosystems between the plant type and characteristics of nutrient uptake. Annual net nitrogen mineralization in soils of different plant communities in the high altitude zone of Spil mountain located in the Mediterranean phytogeographical region of Turkey was investigated throughout one year by field incubation method. Seasonal fluctuations resulting from field incubation were markedly higher in autumn and spring than summer. These are mainly associated with the changes in soil moisture being at minimum in the Mediterranean summer. A significant correlation was developed between the net Nitrate (kg NO3(-)-N ha week(-1)) production and soil water content (p<0.05; r = 0.316 in soil of 0-5 cm; r = 0.312 in soil of 5-15 cm). The results showed that the annual productivity of nitrogen mineralization shows different values depending on communities. Annual net ammonium (NH4(+)-N) production in the soils of each community was negatively estimated. However annual net nitrate (NO3(-)-N) production (0-15 cm) was higher in grassland (27.8 kg ha y(-1)) and shrub (25.0 kg ha y(-1)) than forest (12.4 kg ha y(-1)) community. While annual net N(min) values were close to each other in grassland (14.5 kg ha y(-1)) and shrub (14.1 kg ha y(-1)), but negative in forest community (-3.6 kg ha y(-1)). The reasons for these differences are discussed.     

紫外线B对海洋生态系统的影响

近年来随着臭氧损失现象日益严重，到达地球表面的紫外线B辐射不断加强，海洋生态系统受到了前所未有的威胁。UV-B的增强改变了生物体赖以生存的水体环境，影响了浮游植物对营养盐的吸收同化，抑制了浮游植物的光合作用，从而减少了海洋对CO2的吸收能力，其结果直接增加了温室效应。同时UV-B也直接影响生态系统食物链的各个营养级，伤害海洋病毒、细菌和浮游植物，降低浮游动物和仔稚鱼的成活率，进而影响到整个海洋生态系统的结构和功能。     

Isolation of Oscillatoria spongeliae,the filamentous cyanobacterial symbiont of the marine sponge Dysidea herbacea

The tropical marine sponge Dysidea herbacea (Keller) (Dictyoceratidae: Dysideidae) is always found associated with the filamentous cyanobacterium (blue-green alga) Oscillatoria spongeliae (Schulze) Hauck (Cyanophyceae: Oscillatoriaceae), which occurs abundantly throughout the sponge mesohyl. Intact, metabolically active, trichomes of O. spongeliae were isolated from the sponge by chopping the sponge tissue with a razor blade and squeezing the trichomes into a seawater-based medium containing polyvinylpyrrolidone, bovine serum albumin, dithiothreitol, glycerol, KCl and Na₂CO₃. The isolated cyanobacteria were concentrated by centrifugation and then washed several times in fresh medium. The isolated O. spongeliae have photosynthetic rates which are similar to the intact sponge-alga association for periods of at least 6 h after isolation. Addition of sponge homogenate to the isolated cyanobacteria causes rapid cell lysis.     

Sensory cues associated with host detection in a marine parasitic isopod

Parasite population dynamics and the evolution of life history characteristics are strongly associated with the process of host infection. Parasites with free-living life stages have a narrow window to infect a host and have evolved a number of mechanisms to detect and locate a host. Cymothoa excisa is a parasitic isopod that is commonly found on the Atlantic croaker, Micropogonias undulatus, in the Gulf of Mexico. Here, we determined the infection window that constrains host-searching behavior in C. excisa and tested the behavioral response of the free-swimming larvae (mancae) to visual and chemical cues. Mancae were found to have an infection window of 7 days. Mancae were responsive to both visual and chemical cues. Our findings are the first to show that cymothoid isopods use visual and chemical cues to locate a host and that individuals display a host-locating strategy that maximizes host encounter rate, while reducing energy expenditure.     

Irradiance and temperature regulation of oxygenic photosynthesis and O2 consumption in a hypersaline cyanobacterial mat (Solar Lake, Egypt)

 Short-term effects of temperature and irradiance on oxygenic photosynthesis and O₂ consumption in a hypersaline cyanobacterial mat were investigated with O₂ microsensors in a laboratory. The effect of temperature on O₂ fluxes across the mat–water interface was studied in the dark and at a saturating high surface irradiance (2162 μmol photons m⁻² s⁻¹) in the temperature range from 15 to 45 °C. Areal rates of dark O₂ consumption increased almost linearly with temperature. The apparent activation energy of 18 kJ mol⁻¹ and the corresponding Q ₁₀ value (25 to 35 °C) of 1.3 indicated a relative low temperature dependence of dark O₂ consumption due to mass transfer limitations imposed by the diffusive boundary layer at all temperatures. Areal rates of net photosynthesis increased with temperature up to 40 °C and exhibited a Q ₁₀ value (20 to 30 °C) of 2.8. Both O₂ dynamics and rates of gross photosynthesis at the mat surface increased with temperature up to 40 °C, with the most pronounced increase of gross photosynthesis at the mat surface between 25 and 35 °C (Q ₁₀ of 3.1). In another mat sample, measurements at increasing surface irradiances (0 to 2319 μmol photons m⁻² s⁻¹) were performed at 25, 33 (the in situ temperature) and 40 °C. At all temperatures, areal rates of gross photosynthesis saturated with no significant reduction due to photoinhibition at high irradiances. The initial slope and the onset of saturation (E _k = 148 to 185 μmol photons m⁻² s⁻¹) estimated from P versus E _d curves showed no clear trend with temperature, while maximal photosynthesis increased with temperature. Gross photosynthesis was stimulated by temperature at each irradiance except at the lowest irradiance of 54 μmol photons m⁻² s⁻¹, where oxygenic gross photosynthesis and also the thickness of the photic zone was significantly reduced at 40 °C. The compensation irradiance increased with temperature, from 32 μmol photons m⁻² s⁻¹ at 25 °C to 77 μmol photons m⁻² s⁻¹ at 40 °C, due to increased rates of O₂ consumption relative to gross photosynthesis. Areal rates of O₂ consumption in the illuminated mat were higher than dark O₂ consumption at corresponding temperatures, due to an increasing O₂ consumption in the photic zone with increasing irradiance. Both light and temperature enhanced the internal O₂ cycling within hypersaline cyanobacterial mats. Received: 30 November 1999 / Accepted: 11 April 2000