Sensitivity of Heterogeneous Marine Benthic Habitats to Subtle Stressors

It is important to understand the consequences of low level disturbances on the functioning of ecological communities because of the pervasiveness and frequency of this type of environmental change. In this study we investigated the response of a heterogeneous, subtidal, soft-sediment habitat to small experimental additions of organic matter and calcium carbonate to examine the sensitivity of benthic ecosystem functioning to changes in sediment characteristics that relate to the environmental threats of coastal eutrophication and ocean acidification. Our results documented significant changes between key biogeochemical and sedimentary variables such as gross primary production, ammonium uptake and dissolved reactive phosphorus flux following treatment additions. Moreover, the application of treatments affected relationships between macrofauna communities, sediment characteristics (e.g., chlorophyll a content) and biogeochemical processes (oxygen and nutrient fluxes). In this experiment organic matter and calcium carbonate showed persistent opposing effects on sedimentary processes, and we demonstrated that highly heterogeneous sediment habitats can be surprisingly sensitive to subtle perturbations. Our results have important biological implications in a world with relentless anthropogenic inputs of atmospheric CO₂ and nutrients in coastal waters.     

Estimation of bottom ammonium affinity in the New Caledonia lagoon

Ammonium affinity of New Caledonia lagoon benthic communities was measured during the course of 33 in situ enrichment experiments, in order to estimate the contribution of benthos to ammonium fluxes. Ammonium chloride was injected into enclosures pushed into the sediment, in order to obtain a concentration of 20–22 mol l^-1 in the enclosed water which approximated the interstitial water content. Ammonium kinetic uptake was then followed for two hours. Grey-sand bottom displayed the highest affinity for ammonium, but white-sand and muddy bottom affinity was of the same order of magnitude. Macrophytes, and microphytes (when macrophytes are absent), account for the bulk of ammonium bottom uptake. As a result, grey-sand bottoms with their dense macrophyte cover represent a sink for water column nitrogen and play a key role in nutrient cycling of the lagoon. Correspondence to: G. Boucher     

Benthic Primary Production Budget of a Caribbean Reef Lagoon (Puerto Morelos,Mexico)

High photosynthetic benthic primary production (P) represents a key ecosystem service provided by tropical coral reef systems. However, benthic P budgets of specific ecosystem compartments such as macrophyte-dominated reef lagoons are still scarce. To address this, we quantified individual and lagoon-wide net (P_n) and gross (P_g) primary production by all dominant functional groups of benthic primary producers in a typical macrophyte-dominated Caribbean reef lagoon near Puerto Morelos (Mexico) via measurement of O₂ fluxes in incubation experiments. The photosynthetically active 3D lagoon surface area was quantified using conversion factors to allow extrapolation to lagoon-wide P budgets. Findings revealed that lagoon 2D benthic cover was primarily composed of sand-associated microphytobenthos (40%), seagrasses (29%) and macroalgae (27%), while seagrasses dominated the lagoon 3D surface area (84%). Individual P_g was highest for macroalgae and scleractinian corals (87 and 86 mmol O₂ m⁻² specimen area d⁻¹, respectively), however seagrasses contributed highest (59%) to the lagoon-wide P_g. Macroalgae exhibited highest individual P_n rates, but seagrasses generated the largest fraction (51%) of lagoon-wide P_n. Individual R was highest for scleractinian corals and macroalgae, whereas seagrasses again provided the major lagoon-wide share (68%). These findings characterise the investigated lagoon as a net autotrophic coral reef ecosystem compartment revealing similar P compared to other macrophyte-dominated coastal environments such as seagrass meadows and macroalgae beds. Further, high lagoon-wide P (P_g: 488 and P_n: 181 mmol O₂ m⁻² lagoon area d⁻¹) and overall P_g:R (1.6) indicate substantial benthic excess production within the Puerto Morelos reef lagoon and suggest the export of newly synthesised organic matter to surrounding ecosystems.     

Water column dissolved silica concentration limits microphytobenthic primary production in intertidal sediments

Primary production of microphytobenthos (MPB) contributes significantly to the total production in shallow coastal environments. MPB is a diverse community in which diatoms are usually the main microalgal group. Diatoms require N, P, and other nutrients as with other autotrophs, but in addition require silicate to create their outer cell wall. Therefore, dissolved silica (DSi) might be a potential limiting factor for benthic primary production in areas with reduced freshwater input. To test this hypothesis, a microcosm experiment was conducted using intact sediment cores collected from an intertidal mudflat in the Bay of Cádiz and supplied with increasing concentrations of DSi (0, 5, 10, 25, and 45 μmol · L^?1). After 7 d of enrichment, we determined chlorophyll a and c (Chl a, c) contents, metabolic rates (Net [P_n] and Areal Gross [P_g^A] Production and Light [R_L] and Dark [R_D] Respiration), as well as fluxes of inorganic nutrients across the sediment‐water interface. Chl a and c contents increased significantly with respect to the initial conditions but no differences between treatments were found. Both P_n and P_g^A showed a saturating‐like pattern with silicate concentration, reaching maximum rates at a DSi concentration of 45 μmol · L^?1. The addition of DSi also resulted in an increase of DSi and ammonium uptake by the sediment, which was significantly higher in light than in darkness. Our results clearly show that water column DSi concentrations have a direct impact on benthic primary production, also controlling other related processes such as inorganic nutrient fluxes.     

A revision of the genus Neoneuromus in China (Megaloptera: Corydalidae)

Lake Illawarra, is a typical shallow intertidal coastal barrier lagoon in New South Wales, Australia. This paper reports the first examination of photosynthetic characteristics of benthic microalgae and seagrass in this lake by measuring the oxygen exchange procedure (flux) using sediment-core incubations in the laboratory. Photosynthesis vs irradiance relationships (P–I curves) were generated from measurements made at nine irradiances for microphytobenthos (MPB) at five different water depths (sites) during September 2002. Maximum benthic gross primary production (GP_max) for MPB in this lake tended to decrease with the increasing water depth from Site 5 (about 0.2 m deep) to Site 1 (about 3.0 m deep), and was correlated with decreasing surface sediment Chl-a concentrations.     

Primary Production in a Subtropical Stratified Coastal Lagoon—Contribution of Anoxygenic Phototrophic Bacteria

Anaerobic anoxygenic phototrophic bacteria can be found in the suboxic waters of shallow stratified coastal systems, and may play important roles in the total primary production of subtropical stratified coastal lagoons. We investigated the spatiotemporal variability of light CO₂ fixation and net oxygen production in the stratified Conceição Lagoon (Brazil) in summer and fall of 2007, as well as the contribution of bacteriochlorophyll a (BChl a)-containing bacteria to photosynthetically driven electron transfer. Both chlorophyll a (Chl a) and BChl a varied in space, while only BChl a varied in time (three-fold increase from summer to fall). In summer, net oxygen production and light CO₂ fixation were correlated, with both having higher rates with higher Chl a concentrations in the enclosed region of the lagoon. In fall, CO₂ fixation was decoupled from oxygen production. Denaturing gradient gel electrophoresis revealed that bacterial communities of oxic site 12 and suboxic site 33 formed one cluster, different from other oxic samples within the lagoon. In addition, BChl a/Chl a ratios at these sites were high, 40% and 45%, respectively. Light acted as the main factor controlling the BChl a concentration and CO₂ fixation rates. High turbidity within the enclosed area of the lagoon explained high BChl a and decoupling between CO₂ fixation and oxygen production in oxygenated waters. Contribution of purple sulfur bacteria to total bacterial density in suboxic waters was 1.2%, and their biomass contributed to a much higher percentage (12.2%) due to their large biovolume. Our results indicate a significant contribution of anaerobic anoxygenic bacteria to the primary production of the “dead zone” of Conceição Lagoon.     

Benthic fauna bio-irrigation effects on nutrient regeneration in fish farm sediments

Impacts of organic enrichment and a modified benthic fauna community (caused by fish farming) on benthic mineralization rates and nutrient cycling were studied in sediments at one Danish and one Cypriote fish farm. Sediment organic matter concentration and macrofauna community composition were manipulated in microcosms and changes in total benthic metabolism (oxygen consumption, TCO₂ production), anaerobic metabolism (sulfate reduction rates), nutrient fluxes and sediment parameters were followed for a period of 3 weeks. Mineralization rates were found to be highly correlated with irrigation velocities and largest fauna effects were found in the Danish sediments with the large and active irrigating climax species (Nereis diversicolor and Macoma balthica). Eastern Mediterranean climax species (Glycera rouxii and Naineris laevigata) also stimulated mineralization rates but to a smaller extent due to lower irrigation, whereas the opportunistic species (Capitella in Danish sediment and Hermodice carunculata in Cypriote sediment) showed less effect on mineralization. Ammonium and phosphate release increased with increasing irrigation velocities, but much less in Cyprus indicating higher nutrient retention at the ultra-oligotrophic location compared to eutrophic Danish site. Irrigation velocities, and thus mineralization rates, increased by organic matter loading, indicating larger fauna-induced oxidation in enriched environments. The result implies that a change in fauna structure in fish farm sediment towards smaller opportunistic polychaete species with lower irrigation will result in slower mineralization rates and potentially increase accumulation of organic waste products.     

Microphytobenthos and chironomid larvae attenuate nutrient recycling in shallow‐water sediments

相似文献   

Impact of acidification and eutrophication on macrophyte communities in soft waters. II. Experimental studies

In The Netherlands, there has been a dramatic decline during the last 30 years in the number of stands belonging to the phytosociological alliance Littorellion. Generally, the communities classified within this alliance occur in poorly buffered, oligotrophic waters, with very low phosphate, nitrogen and carbon dioxide levels in the water layer and considerably higher nutrient levels in the sediment. The plant species dominating these communities are isoetids such as Litoorella uniflora (L.) Aschers., Lobelia dortmanna L. and Isoetes lacustris L., which show various adaptations to make successful growth possible under these conditions.Field observations showed that the water where Littorella uniflora had disappeared or strongly decreased could be divided into two groups. A major group (77%) was characterized by the presence of submerged Juncus bulbosus L. and/or Sphagnum species. These water appeared to be strongly acidified (pH < 4.5) and had increased nitrogen levels with ammonium as the dominant N-source. Within this group, the waters with luxuriant growth of Juncus bulbosus and/or Sphagnum spp. had strongly increased carbon dioxide levels in both sediment and water.Different types of experiments proved causal relationships between the observed changes in macrophytes and the changed physico-chemical parameters. Ecophysiological experiments showed that Juncus bulbosus lacks the typical adaptations of the isoetid plant species, i.e. it uses very low amounts of sediment-CO₂ and releases only a little oxygen from the roots. However, Juncus bulbosus is more able than Littorella uniflora to use CO₂ from the water layer. From the nutrient-uptake experiments, the decreased nitrate and increased ammonium levels seem to be favourable to Juncus bulbosus. The culture experiments clearly demonstrated that the biomass of Juncus bulbosus only increased strongly when the sediment was poorly buffered and the pH of water was low. When combining factors like CO₂ enrichment of the sediment, with and without phosphate, and/or ammonium enrichment of the water in the culture experiments, it is clearly shown that phosphate and/or ammonium enrichment without CO₂ enrichment do not lead to an increase in biomass of Juncus bulbosus. Therefore, it is obvious that the changes in the macrophyte community can be ascribed primarily to changes in the carbon budget as a result of acidification.A minor group of waters (23%) was characterized by the absence of submerged Juncus bulbosus and/or Sphagnum spp. In most of these waters, submerged plant species occurred, such as Myriophyllum alterniflorum DC or non-rooted species such as Riccia fluitans L. These waters were not acidified, and generally had an increased alkalinity and higher nitrogen and phosphate levels of sediment and/or water. Culture experiments showed that phosphate enrichment of the sediment alone leads to luxuriant growth of submerged macrophyte species such as Myriophyllum alterniflorum, whereas phosphate enrichment of both sediment and water leads to mass development of non-rooted plant species such as Riccia fluitans.     

Population Ecology of Nitrifiers in a Stream Receiving Geothermal Inputs of Ammonium

The distribution, activity, and generic diversity of nitrifying bacteria in a stream receiving geothermal inputs of ammonium were studied. The high estimated rates of benthic nitrate flux (33 to 75 mg of N · m⁻² · h⁻¹) were a result of the activity of nitrifiers located in the sediment. Nitrifying potentials and ammonium oxidizer most probable numbers in the sediments were at least one order of magnitude higher than those in the waters. Nitrifiers in the oxygenated surface (0 to 2 cm) sediments were limited by suboptimal temperature, pH, and substrate level. Nitrifiers in deep (nonsurface) oxygenated sediments did not contribute significantly to the changes measured in the levels of inorganic nitrogen species in the overlying waters and presumably derived their ammonium supply from ammonification within the sediment. Ammonium-oxidizing isolates obtained by a most-probable number nonenrichment procedure were species of either Nitrosospira or Nitrosomonas, whereas all those obtained by an enrichment procedure (i.e., selective culture) were Nitrosomonas spp. The efficiency of the most-probable-number method for enumerating ammonium oxidizers was calculated to be between 0.05 and 2.0%, suggesting that measurements of nitrifying potentials provide a better estimate of nitrifying populations.     

The control of nitrate and ammonium concentrations in a coral reef lagoon

One Tree Reef lagoon is surrounded by an emergent rim which restricts exchange between lagoonal waters and the surrounding ocean. For this reason, the loss rate of dissolved inorganic nitrogen (DIN) through mixing processes is slow in the central lagoon compared to rates of advective input, uptake, regeneration and loss to the atmosphere. We present some hypotheses concerning the importance of these fluxes to the observed patterns of concentration of nitrate+nitrite and ammonium. A scaling analysis of these fluxes indicates that the relative influence of advection across the windward reef crest on lagoonal concentrations changes with season and differs for the two forms of DIN. Advective flux, dominated by DIN derived from production on the algal pavements of the reef crest, is significant in controlling DIN concentration in the peripheral regions of the lagoon. Loss to the atmosphere is a more important flux from the nitrate+nitrite pool in the centre of the lagoon, particularly in summer. Regeneration is a significant input to the ammonium pool of the central lagoon in winter. The relative magnitudes of all fluxes are more similar to each other in the summer than the winter, indicating the potential for shifts in the dominance hierarchy at small time and space scales. One form of DIN in One Tree Reef lagoonal waters (nitrate+nitrite) is controlled by input and another (ammonium) by recycling as well as input. The relative importance of these fluxes changes as a result of temperature pertubations at the physiological level as well as the rate of water turnover at the system level. It is proposed that the degree of consistency of the seasonal concentration patterns is a function of the period, rather than the amplitude of the temporal oscillations in the fluxes controlling these concentrations. This has important implications for sampling strategies. This paper provides a conceptual framework for hypothesis testing at a manageable scale, in the context of ecosystem function.     

49 Use of biomarkers and core incubations to investigate how changes in benthic ecosystems influence sediment n fluxes in western australia

The west coast of Australia is dominated by warm, oligotrophic waters, shallow carbonate-rich sediments and little terrigeneous nutrient input. Irradiance and light penetration are generally high so that benthic primary production is substantial and can equal that in the water column. Microphytobenthos has been shown to have a major influence on nutrient cycling in northern hemisphere waters, but little work has been published on this subject describing the Western Australian marine environment. This presentation focuses upon anthropogenic impacts on microphytobenthos and sediment-water column nitrogen cycling. The effects of artificial enclosures (harbours) on microphytobenthic communities and nitrogen cycling were examined over an annual cycle at 2 locations in Cockburn Sound, Western Australia. Core and bulk sediment samples were taken from bare sandy areas in water 6–7 m deep at paired sites inside and outside sea walls. Ex-situ core incubations were used to determine rates of N2 fixation, denitrification, fluxes of macronutrients and oxygen. Bulk sediment samples were analysed for pigments, fatty acids, sterols and grain size. Significant differences were found in both flux and biomarker data between the 2 areas, inside and outside the harbours and between summer and winter. Biomarker data gave information about the algal, bacterial and faunal composition of the sediments and how it changed across the same temporal and spatial scales as the fluxes. The combination of process studies and biomarker information promises to be a powerful tool for resolving a range of questions on the magnitude and mechanisms of nitrogen inputs and exports from these shallow ecosystems.     

Carbon and Nitrogen Cycling in a Shallow Productive Sub-Tropical Coastal Embayment (Western Moreton Bay, Australia): The Importance of Pelagic–Benthic Coupling

Climatic variables, water quality, benthic fluxes, sediment properties, and infauna were measured six times over an annual cycle in a shallow sub-tropical embayment to characterize carbon and nutrient cycling, and elucidate the role of pelagic–benthic coupling. Organic carbon (OC) inputs to the bay are dominated by phytoplankton (mean 74%), followed by catchment inputs (15%), and benthic microalgae (BMA; 9%). The importance of catchment inputs was highly variable and dependent on antecedent rainfall, with significant storage of allochthonous OC in sediments following high flow events and remineralization of this material supporting productivity during the subsequent period. Outputs were dominated by benthic mineralization (mean 59% of total inputs), followed by pelagic mineralization (16%), burial (1%), and assimilation in macrofaunal biomass (2%). The net ecosystem metabolism (NEM = production minus respiration) varied between ?4 and 33% (mean 9%) of total primary production, whereas the productivity/respiration (p/r) ranged between 0.96 and 1.5 (mean 1.13). Up to 100% of the NEM is potentially removed via the demersal detritivore pathway. Dissolved inorganic nitrogen (DIN) inputs from the catchment contributed less than 1% of the total phytoplankton demand, implicating internal DIN recycling (pelagic 23% and benthic 19%) and potentially benthic dissolved organic nitrogen (DON) fluxes (27%) or N fixation (up to 47%) as important processes sustaining productivity. Although phytoplankton dominated OC inputs in this system, BMA exerted strong seasonal controls over benthic DIN fluxes, limiting pelagic productivity when mixing/photic depth approached 1.3. The results of this study suggest low DIN:TOC and net autotrophic NEM may be a significant feature of shallow sub-tropical systems where the mixing/photic depth is consistently less than 4.     

Effect of nutrient pulses on photosynthesis of Chaetomorpha linum from a shallow Mediterranean coastal lagoon

The influence of nitrogen and phosphorus pulses on Chaetomorpha linum (Muller) Kutzing growth and photosynthesis was studied in laboratory experiments. Photosynthesis and growth of C. linum from Tancada lagoon seems limited by both nitrogen and phosphorus, as indicated by the high rate (4.7–11.6 mg O₂ g⁻¹ dry weight h⁻¹) of light-saturated photosynthesis (P_m) and growth rates observed under nitrogen plus phosphorus enrichment in relation to enrichment by nitrogen alone (2.9–7.6 mg O₂ g⁻¹ dry weight h⁻¹). Significant increase in nitrogen and phosphorus content as percentage of dry weight was observed in C. linum fertilized with a single nutrient or with nitrogen plus phosphorus. In Tancada lagoon, when availability of nitrogen to primary producers is by pulses, an increase of nitrate concentration in the water column (from 6 to 100 μM) has a greater effect on growth of C. linum (growth rate: 0.13 day⁻¹) than an increase in ammonium concentration (from 20 to 100 μM and growth rate: 0.11 day⁻¹). For a given thallus nitrogen content (0.6–1.4% N), both P_m and the photosynthetic efficiency (α) normalized to dry weight were correlated (r² = 0.73, p < 0.005) indicating that variations in electron transport were coupled to variations in C-fixation capacity. Optimizing both α and P_m may be a general characteristic of thin-structured opportunistic algae in more variable estuarine environments.     

Benthic phosphorus regeneration in the Potomac River Estuary

The flux of dissolved reactive phosphate from Potomac riverine and estuarine sediments is controlled by processes occurring at the water-sediment interface and within surficial sediment.In situ benthic fluxes (0.1 to 2.0 mmoles m⁻² day⁻¹) are generally five to ten times higher than calculated diffusive fluxes (0.020 to 0.30 mmoles m⁻² day⁻¹). The discrepancy between the two flux estimates is greatest in the transition zone (river mile 50 to 70) and is attributd to macrofaunal irrigation. Bothin situ and diffusive fluxes of dissolved reactive phosphate from Potomac tidal river sediments are low while those from anoxic lower estuarine sediments are high. The net accumulation rate of phosphorus in benthic sediment exhibits an inverse pattern. Thus a large fraction of phosphorus is retained by Potomac tidal river sediments, which contain a surficial oxidized layer and oligochaete worms tolerant of low oxygen conditions, and a large fraction of phosphorus is released from anoxic lower estuary sediments. Tidal river sediment pore waters are in equilibrium with amorphous Fe (OH)₃ while lower estuary pore waters are significantly undersaturated with respect to this phase. Benthic regeneration of dissolved reactive phosphorus is sufficient to supply all the phosphorus requirements for net primary production in the lower tidal river and transition-zone waters of the Potomac River Estuary. Benthic regeneration supplies approximately 25% as much phosphorus as inputs from sewage treatment plants and 10% of all phosphorus inputs to the tidal Potomac River. When all available point source phosphorus data are put into a steady-state conservation of mass model and reasonable coefficients for uptake of dissolved phosphorus, remineralization of particulate phosphorus, and sedimentation of particulate phosphorus are used in the model, a reasonably accurate simulation of dissolved and particulate phosphorus in the water column is obtained for the summer of 1980.     

Bacterial and archaeal phylogenetic diversity associated with swine sludge from an anaerobic treatment lagoon

Over the last decades, the demand for pork products has increased significantly, along with concern about suitable waste management. Anaerobic-lagoon fermentation for swine-sludge stabilization is a good strategy, although little is known about the microbial communities in the lagoons. Here, we employed a cloning- and sequencing-based analysis of the 16S rRNA gene to characterize and quantify the prokaryotic community composition in a swine-waste-sludge anaerobic lagoon (SAL). DNA sequence analysis revealed that the SAL library harbored 15 bacterial phyla: Bacteroidetes, Cloroflexi, Proteobacteria, Firmicutes, Deinococcus-Thermus, Synergystetes, Gemmatimonadetes, Chlorobi, Fibrobacteres, Verrucomicrobia and candidates division OP5, OP8, WWE1, KSB1, WS6. The SAL library was generally dominated by carbohydrate-oxidizing bacteria. The archaeal sequences were related to the Crenarchaeota and Euryarchaeota phyla. Crenarchaeota predominated in the library, demonstrating that it is not restricted to high-temperature environments, being also responsible for ammonium oxidation in the anaerobic lagoon. Euryarchaeota sequences were associated with the hydrogenotrophic methanogens (Methanomicrobiales and Methanobacteriales). Quantitative PCR analysis revealed that the number of bacterial cells was at least three orders of magnitude higher than the number of archaeal cells in the SAL. The identified prokaryotic diversity was ecologically significant, particularly the archaeal community of hydrogenotrophic methanogens, which was responsible for methane production in the anaerobic lagoon. This study provided insight into the archaeal involvement in the overall oxidation of organic matter and the production of methane. Therefore, the treatment of swine waste in the sludge anaerobic lagoon could represent a potential inoculum for the start-up of municipal solid-waste digesters.     

Benthic community response to a passive fishing gear in a coastal lagoon (South Brazil)

The influence of a passive shrimp fishing gear on benthic communities was studied at Laguna Estuarine System (South Brazil), a shallow choked coastal lagoon. The gear is composed by a group of fyke nets (25 mm mesh size) set in contact to the bottom, fixed with stakes forming a cage-like structure (around 30 m²). Samplings were conducted in the two main fishery areas of the estuarine system, Mirim (sand bottoms) and Imaruí (muddy bottoms) lagoon, in May 2005. In each area, 10 fyke net enclosures and 10 nearby sites without nets (control) were sampled. Microphytobenthos biomass (chlorophyll a and phaeopigments), number of taxa/species, density, Hill’s number N ₁ and N ₂, and estimated number of species (ES₁₀₀) were used as community attributes. For the nematodes, values of the maturity index and abundance of Wieser’s feeding type were used as well. The effects of the small-scale passive shrimp fishing gear on the coastal lagoon bottoms were dependent on the benthic component analyzed and the type of sediment. Whereas macrofauna was not affected by the net enclosures, meiofauna and nematodes, particularly from the mud sites were. At the sand site, the fyke net enclosures caused a decrease in the microphytobenthos biomass and changed the relative abundances of non-selective deposit feeding and epigrowth-feeding nematodes. The results indicated that small-scale static nets, such as the studied fyke enclosures, produced low intensity levels of disturbance. However, the enclosed area by nets at Laguna had already reached around 25,000 m². Given the large proportion of the coastal population involved and the area closed by nets, management policies should consider site-specific differences within the same estuarine system.     

Microphytobenthos production in the Gulf of Fos, French Mediterranean coast

Microphytobenthic oxygen production was studied in the Gulf of Fos (French Mediterranean coast) during 1991/1992 using transparent and dark benthic chambers. Nine stations were chosen in depths ranging from 0.5 to 13 m, which represents more than 60% of bottoms in the Gulf. Positive net microphytobenthic oxygen production was seasonally detected down to 13 m; the maximum value attained was 60 mg O₂ m⁻² h⁻¹ (0.7–0.8 g O₂ m⁻² d⁻¹) in sediments at 0.5 m depth during spring and winter. Respiration rates were maximum in the sediments located at the mussel farm (5 m), in the center of the Gulf, with 135 mg O₂ m⁻² h⁻¹ in spring (3.2 g O₂ m⁻² d⁻¹); in the other locations, it ranged from 3.3 to 58.2 mg O₂ m⁻² h⁻¹ (0.08–1.4 g O₂ m⁻² d⁻¹). Compared to phytoplankton, microphytobenthos production was higher only in the bottoms < 1 m depth. In deeper bottom waters, phytoplankton production could be absent due to light limitation, while microphytobenthos was still productive. Phytoplankton production m⁻² was generally higher than microphytobenthic production. Microphytobenthic biomass, higher than phytoplanktonic, varied from 27 to 379 mg Chl a m⁻², the maximum in the mussel farm sediments, with the minimum in sandy shallow bottoms. Pigment analysis showed that microphytobenthos consisted mainly of diatoms (Chl c and fucoxanthin) but other algal groups containing Chl b could become seasonally important. A Principal Component Analysis suggested that the main statistical factors explaining the distribution of our observations may be interpreted in terms of enrichment in phaeopigments and light; the role of Chl a appearing paradoxically as secondary in benthic production rates. Phaeopigments are mainly constituted by phaeophorbides, which indicate grazing processes. The influence of the mussel farm on the oxygen balance is noticeable in the whole Gulf.     

BENTHIC AND PLANKTONIC ALGAL COMMUNITIES IN A HIGH ARCTIC LAKE: PIGMENT STRUCTURE AND CONTRASTING RESPONSES TO NUTRIENT ENRICHMENT1

We investigated the fine pigment structure and composition of phytoplankton and benthic cyanobacterial mats in Ward Hunt Lake at the northern limit of High Arctic Canada and the responses of these two communities to in situ nutrient enrichment. The HPLC analyses showed that more than 98% of the total pigment stocks occurred in the benthos. The phytoplankton contained Chrysophyceae, low concentrations of other protists and Cyanobacteria (notably picocyanobacteria), and the accessory pigments chl c₂, fucoxanthin, diadinoxanthin, violaxanthin, and zeaxanthin. The benthic community contained the accessory pigments chl b, chl c₂, and a set of carotenoids dominated by glycosidic xanthophylls, characteristic of filamentous cyanobacteria. The black surface layer of the mats was rich in the UV‐screening compounds scytonemin, red scytonemin‐like, and mycosporine‐like amino acids, and the blue‐green basal stratum contained high concentrations of light‐harvesting pigments. In a first bioassay of the benthic mats, there was no significant photosynthetic or growth response to inorganic carbon or full nutrient enrichment over 15 days. This bioassay was repeated with increased replication and HPLC analysis in a subsequent season, and the results confirmed the lack of significant response to added nutrients. In contrast, the phytoplankton in samples from the overlying water column responded strongly to enrichment, and chl a biomass increased by a factor of 19.2 over 2 weeks. These results underscore the divergent ecophysiology of benthic versus planktonic communities in extreme latitudes and show that cold lake ecosystems can be dominated by benthic phototrophs that are nutrient sufficient despite their ultraoligotrophic overlying waters.     

Reevaluation of the nutrient mineralization process by infaunal bivalves (Ruditapes philippinarum) in a shallow lagoon in Hokkaido, Japan

Previous estimations of nutrient mineralization in the water column by infaunal bivalves might have been overestimated because of underestimation of the uptake process by microphytobenthos in the field. We conducted field surveys of environmental conditions and quantitative sampling of Ruditapes philippinarum in a shallow lagoon system (Hichirippu Lagoon, eastern Hokkaido, Japan) in August 2006. We recorded the spatial distribution pattern and the molar ratio of dissolved inorganic nutrients to determine the limiting nutrients for microphytobenthos, to evaluate the input and output of nutrients at the entrance of the lagoon station, and to estimate potential nutrient mineralization by R. philippinarum. Our aim was to reevaluate the nutrient mineralization process by infaunal bivalve species. In this study, the mean standing stock of microphytobenthos inhabiting surface sediment (5 mm thick) on the tidal flats was 100 times higher than that of phytoplankton (1 m depth). Low N/P and high Si/N ratios (mean = 2.6 and 17.6, respectively) near the entrance of the lagoon compared to those of microphytobenthos (N:P:Si = 10.1:1:18) clearly suggest N deficiency. The flux of NH₄-N coming into the lagoon was 3.4 kmolN d^− 1, and the flux out was − 3.7 kmolN d^− 1. Thus, assuming that there would have been no phytoplankton and microphytobenthos uptake during the day, 0.3 kmolN d^− 1 of NH₄-N was produced within the lagoon. However, the NH₄-N mineralization rate of the clams has been estimated to be approximately 7.7 ± 6.8 kmolN d^− 1. Thus, 96% (7.4 kmolN d^− 1, i.e., 7.7 kmolN d^− 1 minus 0.3 kmolN d^− 1) of the NH₄-N mineralized by the clam was consumed by microphytobenthos. In contrast, if all the NH₄-N inflow (3.1 kmolN d^− 1) was consumed by the microalgae before outflow, 52% (4.0 kmolN d^− 1, i.e., 7.7 kmolN d^− 1 minus 3.7 kmolN d^− 1) of the NH₄-N mineralized by the clams should have been consumed by microphytobenthos. Microphytobenthos on the tidal flats (11.3 ± 11.8 kmolN) used all of the surplus nutrients (between 4.0 and 7.4 kmolN d^− 1), and the temporal division rate [=(NH₄-N uptake)/(standing stock of microphytobenthos)] of microphytobenthos would have to be between 0.35 and 0.65 d^− 1. Residual NH₄-N (0.3 - 3.7 kmolN d^− 1) was the water-column source and accounted for 12-148% of NH₄-N in the water column near the entrance of the lagoon (2.5 ± 1.4 kmolN) per day. This is the first field-based observation with a quantitative evaluation of nutrient mineralization by infaunal bivalves and nutrient uptake by microphytobenthos.