It's a dirty job but someone has to do it: The role of marine benthic macrofauna in organic matter turnover and nutrient recycling to the water column

Benthic macrofaunal populations through their feeding, bioturbation, burrow construction and sediment irrigation activities have profound influences on organic matter inputs to marine sediments (biodeposition) and on the vertical distribution of deposited organic matter within the sediment. These effects in turn influence the rates and pathways of organic matter mineralisation, and element cycles. Similarly, bioturbation, burrow construction and burrow irrigation are major determinants of sediment-water column fluxes of oxygen and nutrients.

In this review, I discuss the influences of the different benthic macrofaunal feeding (functional) groups on mineralisation processes and sediment-water column fluxes of particulate and dissolved nutrients. How these effects influence diagenic processes, the balance between aerobic and anaerobic processes, and the redox status of the surficial sediments. Finally, I discuss some of the limitations of the predominantly laboratory techniques which have been used to study “macrofaunal effects” and how this hinders the inclusion of the effects in quantitative sediment biogeochemical models.     

Microbial reaction rates and bacterial communities in sediment surrounding burrows of two nereidid polychaetes (Nereis diversicolor and N. virens)

The effects of infaunal mode of life on sediment properties, microbial reaction rates, as well as abundance and composition of bacterial communities were studied in sediment surrounding burrows (mucus lining, oxidised wall, ambient anoxic and surface sediment) of two closely related, but behaviourally different, nereidid polychaete worms: the facultative suspension-feeder Nereis (Hediste) diversicolor and the obligate deposit-feeder Nereis (Neanthes) virens. Burrow sediment of the two species was collected from two adjacent (50 m distance) shallow sandy locations (Kertinge Nor, Denmark). The burrow lining and wall of both polychaete species were enriched in organic matter originating from mucous secretions by the inhabitants and phytoplankton trapped through irrigation. This was more evident for N. diversicolor that shows a significantly higher irrigation rate than N. virens. Both the organic matter mineralisation rates (based on anaerobic incubations) and bacterial abundance were higher along the burrow linings and walls. However, accumulation of porewater TCO₂ and dissolved organic carbon in sediments adjacent to burrows increased most rapidly in the presence of N. diversicolor, suggesting higher heterotrophic activity associated with this species. Surprisingly, bacterial abundance was lower around burrows of N. diversicolor than those from N. virens indicating that burrow environments from the first species harbour a more active bacterial community. Molecular fingerprints of the 16S rRNA gene from bacterial communities showed that the composition of the burrow linings and walls resembled the ambient anoxic sediment rather than the oxic sediment surface. On the other hand, the bacterial fingerprints of the sediment surrounding the burrows of the two polychaete species were markedly different suggesting either a site-specific difference in sediment parameters or a significant species-specific impact of the burrow inhabitants.     

Microbial activity and accumulation of organic matter in the burrow of the mud shrimp, <Emphasis Type="Italic">Upogebia major</Emphasis> (Crustacea: Thalassinidea)

Microbial activity and accumulation of organic matter in the burrow of the thalassinidean mud shrimp, Upogebia major, were studied on a tidal flat along the northern coast of Tokyo Bay, Japan. The burrow of U. major is Y-shaped with an upper U-shaped part plus a lower I-shaped part. Its lower part can extend to a depth of 2 m. In the present study, we compare electron transport system activity (ETSA), bacterial abundance and organic matter content [total organic carbon (TOC), total nitrogen (TN) and chlorophyll a (chl. a)] of the burrow wall sediment with the tidal flat surface and non-burrow sediments. We also compared the U- and I-shaped part in terms of these parameters. ETSA in the burrow wall was higher than at the tidal flat surface in the warmer season, and was always higher than at surrounding non-burrow sediments. Bacterial abundance in the burrow wall was higher than at the tidal flat surface and surrounding sediment. TOC and TN contents in the burrow wall were two to three times higher than those at the tidal flat surface and non-burrow sediments, regardless of season. However, there was no significant difference in chl. a content between burrow wall and tidal flat surface. These results suggest that organic enrichment of the burrow wall is a result of organic matter particles such as phytodebris accumulation along the burrow wall. For all parameters of the burrow walls, no statistical differences were found between the two parts. Present results indicate that U. major actively transports the water containing suspended organic particles not only through the U-part but also into the deeper I-part. These indicate that burrow of the mud shrimp provides a dynamic environment for microbial community in tidal flat sediment.     

Trait-mediated indirect effects of predatory fish on microbial mineralization in aquatic sediments

Sediment-dwelling zoobenthos stimulate the mineralization of organic matter and alter nutrient cycling by aerating the sediment via burrows, by feeding on detritus, and by redistributing particles. Here we experimentally revealed that abundant benthic chironomids (Chironomus riparius) can perceive predatory fish (Rutilus rutilus) via chemical cues (kairomones) and spend less time foraging at the sediment surface and more time hiding in their burrows. This predator avoidance behavior significantly increased the exposure of freshly deposited organic particles to oxygen by reducing their burial to subsurface layers and by enhancing the aeration of subsurface layers via burrow ventilation, conditions that together increased the rate of organic matter mineralization. These results demonstrate that predatory fish can exert trait-mediated effects on benthic communities that in turn alter basic ecosystem processes related to nutrient cycling.     

Modelling of the relationship moisture content to nutrient transformation rate in river sediments

The modelling of the transformation processes in the river sediments is an important and still not investigated problem that is a key for water quality. The relation between nutrients transformation rate and moisture in sediments was investigated by means of analogous modelling. The mathematical dependences of the connection were derived, and only two of them were linear—at the early stage of COD (organic) change and at the late stage of transformation of phosphates that metabolically connected the biodegradation of organics with oxidative phosphorylation. The rest mathematical dependences were the polynomials of third order. This study confirmed that the leading factors which influence the transformation rate of nutrients are the sediment surface, particular organic matter and porosity of the medium, as well as the concentration on the sediment surface of micro-organisms, enzymes, co-metabolites, etc. The presented data from ecological modelling of the transformation processes in the parafluvial sediments showed how can the hydrological models be enlarged and enriched in their application to the key and critical (risk) fluctuations in the river functioning. The mathematical dependences and coefficients were extracted in our complex study to be included in HSPF model as an enlarged module. As far as the investigated transformation processes in the parafluvial have more universal significance, the results can be applied in the other models, bounded river water and sediment quality.     

生物扰动对沉积物中污染物环境行为与生物可利用性的影响

疏水性有机污染物进入环境水体后易于与沉积物结合,对沉积物中的底栖动物造成危害。底栖动物引起的生物扰动作用可以通过改变沉积物的地球化学性质,对其中污染物的赋存形态、迁移转化和生物可利用性产生重要影响。在综述了国内外生物扰动影响沉积物中污染物环境行为和生物可利用性的最新研究进展基础上,重点讨论了沉积物颗粒交换、水体环境条件改变、疏水有机污染物解吸释放过程以及对生物扰动影响的定量化表征。最后对该研究方向进行了展望,指出应重点研究多种污染物及不同生物共存条件下的生态效应,以及造成沉积物扰动的影响因素的定量化表征等。     

Biotic control of stream fluxes: spawning salmon drive nutrient and matter export

Organisms can control movements of nutrients and matter by physically modifying habitat. We examined how an ecosystem engineer, sockeye salmon (Oncorhynchus nerka), influences seasonal fluxes of sediments, nitrogen (N), and phosphorus (P) in streams of southwestern Alaska. The purpose of this study was to investigate whether salmon act as net importers or net exporters of matter and nutrients from streams and how these roles change as a function of salmon population density. We measured discharge and concentrations of suspended sediments and total N and P every 7-14 days for up to four summers in 10 streams spanning a gradient in salmon densities. We statistically allocated whole-season fluxes to salmon activities, such as excretion and bioturbation, and to export by hydrologic discharge. In addition, we used counts of spawning salmon to estimate nutrient and matter imports by salmon to streams. Large seasonal pulses of suspended sediments, P, and N were associated with salmon spawning activities, often increasing export an order of magnitude higher than during pre-salmon levels. Years and streams with more salmon had significantly higher levels of export of sediments and nutrients. In addition, years with higher precipitation had higher background export of P and N. Salmon exported an average of the equivalent of 189%, 60%, and 55% of total matter, P, and N that salmon imported in their bodies. The relative magnitude of export varied; salmon exported more than their bodies imported in 80%, 20%, and 16% across all streams and years for sediments, P, and N, respectively. A bioassay experiment indicated that the P exported by salmon is directly available for use by primary producers in the downstream lake. These results demonstrate that salmon not only move nutrients upstream on large spatial scales via their migration from the ocean and subsequent death, but also redistribute matter and nutrients on finer spatial scales through their spawning activities.     

Mineralization of particulate organic matter derived from coral-reef organisms in reef sediments of the Gulf of Aqaba

In situ and laboratory incubation experiments in a fringing reef in the Gulf of Aqaba were performed to study degradation rates of particulate organic matter in reef sediments. Coral mucus, clam eggs, and zooxanthellae were used as model particulate organic compounds for these experiments. Aerobic and anaerobic mineralization rates were calculated by dissolved inorganic carbon (DIC) and O₂ fluxes from the sediments under different particulate organic matter additions. Fast enhancement (approximately twofold) of O₂ and DIC fluxes were found with the addition of coral mucus and clam eggs compared with control incubations without addition. Most of the degradation is believed to have occurred anaerobically rather than aerobically (DIC:O₂ ratios were 4.3–28.1). Higher degradation rates of coral mucus and clam eggs were estimated in carbonate sediment than in silicate sediment (1.2–1.6-fold), which was attributed to the different physical and chemical properties of both sediments. Our study shows the significance of the reef sediment as a suitable site for microbial degradation of particulate organic material excreted from different reef community organisms. This may increase the regeneration of nutrients in the reef environment necessary to sustain high biological productivity.     

Mineralization of particulate organic matter derived from coral-reef organisms in reef sediments of the Gulf of Aqaba

In situ and laboratory incubation experiments in a fringing reef in the Gulf of Aqaba were performed to study degradation rates of particulate organic matter in reef sediments. Coral mucus, clam eggs, and zooxanthellae were used as model particulate organic compounds for these experiments. Aerobic and anaerobic mineralization rates were calculated by dissolved inorganic carbon (DIC) and O₂ fluxes from the sediments under different particulate organic matter additions. Fast enhancement (approximately twofold) of O₂ and DIC fluxes were found with the addition of coral mucus and clam eggs compared with control incubations without addition. Most of the degradation is believed to have occurred anaerobically rather than aerobically (DIC:O₂ ratios were 4.3-28.1). Higher degradation rates of coral mucus and clam eggs were estimated in carbonate sediment than in silicate sediment (1.2-1.6-fold), which was attributed to the different physical and chemical properties of both sediments. Our study shows the significance of the reef sediment as a suitable site for microbial degradation of particulate organic material excreted from different reef community organisms. This may increase the regeneration of nutrients in the reef environment necessary to sustain high biological productivity.     

Variable feeding behavior in three species ofMacoma (Bivalvia: Tellinacea) as a response to water flow and sediment transport

Many species of tellinacean bivalves mainly deposit-feed but are also known to vary their feeding behavior in response to predation and to the availability of suspended organic matter relative to that of sedimentary organic matter. This study showed that three species of the Pacific genusMacoma (Macoma nasuta Conrad, 1937,Macoma secta Conrad, 1837, andMacoma inquinata Deshayes, 1855, from the San Juan Islands, Washington, USA) varied their deposit-feeding behavior in response to water flow near the sediment-water interface and to sediment transport. In the summers of 1987, 1988 and 1989, water velocity was varied in a large racetrack flume, in order to change both water velocity and bottom sediment transport. In quiet water, the inhalant siphon was protruded far from the siphon hole. As water velocity increased, with little or no sediment transport, the deposit-feeding radius decreased. Qualitative observations suggested that this was related to the drag on the siphon. InM. secta, the distance of siphon protrusion was not related to body size. Under conditions of higher near-bottom water velocity, combined with bottom sediment transport, some individuals ceased to deposit-feed, while others sustained feeding by ingesting sediment within the siphon hole, at least 1 cm beneath the sediment-water interface. These results suggest that hydrodynamic conditions are a major determinant of feeding behavior, and previous explanations of variable feeding behavior as a response to predation may have to be adjusted to accommodate this expanded set of responses.     

影响沉积物-水界面持久性有机污染物迁移行为的因素研究

沉积物-水界面是自然水体在物理、化学和生物特征等方面差异性最显著的环境边界,界面及附近发生的物理和化学反应,如吸附和解吸、迁移和转化、扩散和掩埋以及生物扰动等作用是控制和调节水体和沉积物之间物质输送和交换的重要途径.持久性有机污染物由于其具有致癌、致畸、致突变及内分泌干扰作用而受到人们的广泛关注,它们在沉积物-水界面的...     

The Influence of Anthropogenic Contamination On Metals Release From Coastal Sediment Suspensions with Aerated Sea Water

The zinc, cadmium, copper and lead release from the uncontaminated and contaminated coastal sediments with aerated sea water was studied. the metals transfer to the dissolved forms was monitored during one-two months by differential pulse anodic voltammetry (DPASV). the sediments with different initial degree of contamination were sampled in the Peter the Great Bay, Sea of Japan around Vladivostok-the biggest city in Russian Far East. Sediment contamination by metals led to increased release to solution of zinc due to sulphide oxidation and cadmium on account of organic matter decomposition. the copper behaviour was complicated by strong binding with organic matter and enhancements of copper release can only be seen in sediment with a low organic matter content. Significant lead transfer to dissolved forms was not observed regardless of sediment contamination. the temperature affected the release of cadmium and copper through enhanced organic matter destruction. the dissolved metal fluxes from the sediment transformation are compared with metal fluxes towards bottom. Such comparisons show that second contamination by dissolved metals of the studied coastal environment may be important for cadmium only.     

The Influence of Anthropogenic Contamination On Metals Release From Coastal Sediment Suspensions with Aerated Sea Water

The zinc, cadmium, copper and lead release from the uncontaminated and contaminated coastal sediments with aerated sea water was studied. the metals transfer to the dissolved forms was monitored during one-two months by differential pulse anodic voltammetry (DPASV). the sediments with different initial degree of contamination were sampled in the Peter the Great Bay, Sea of Japan around Vladivostok-the biggest city in Russian Far East. Sediment contamination by metals led to increased release to solution of zinc due to sulphide oxidation and cadmium on account of organic matter decomposition. the copper behaviour was complicated by strong binding with organic matter and enhancements of copper release can only be seen in sediment with a low organic matter content. Significant lead transfer to dissolved forms was not observed regardless of sediment contamination. the temperature affected the release of cadmium and copper through enhanced organic matter destruction. the dissolved metal fluxes from the sediment transformation are compared with metal fluxes towards bottom. Such comparisons show that second contamination by dissolved metals of the studied coastal environment may be important for cadmium only.     

In situ assessment of modification of sediment properties by burrowing invertebrates

Benthic organisms can significantly alter the physical properties of marine sediments, but it has hitherto been difficult to assess and quantify the effects of bioturbation. In situ geophysical techniques offer new methods for measuring these effects: measurement of acoustic shear-wave velocity and electrical resistivity allows nondestructive assessment of the properties of the grain framework and pore-fluid matrix, respectively, of the seabed sediment. The influence of burrowing invertebrates on the structural properties of sandy sediments at intertidal locations on the coast of Wales (UK) was investigated during the periol 1986–1987 using these techniques. Three species (Arenicola marina, Corophium arenarium and Lanice conchilega) were selected on the basis of their contrasting styles of burrow construction. All three species produced measurable and significant, although different, changes in bed properties. They modified shear-wave propagation through the bed by changing bed rigidity: while A. marina and C. arenarium decreased rigidity by creating open burrows, L. conchilega increased rigidity by building shell-lined tubes. All produced a decrease in electrical resistivity by altering porosity and/or tortuosity, which implies an increase in permeability; these changes were attributable not only to the presence of the burrows but also to modification of the between-burrow sediment texture and bed properties.     

Pelagic-benthic coupling in a subtidal system of the North-Western Mediterranean

Suspended particulate matter, zooplankton, and macrobenthos dynamics were investigated in a shallow area of the Ligurian Sea (north-west Mediterranean) characterized by wide temporal variability over an annual cycle. As indicated by multivariate analyses, the seasonal dynamics can be summarized as follows: (1) a late winter-early spring phytoplankton bloom followed by high zooplankton and macrobenthos densities during the spring months; (2) low-quality particulate suspended matter in summer, and an increase in the importance of zooplankton taxa with a wide range of feeding strategies, a decrease in macrofaunal abundance, and an increase in deposit-feeders and predators; and (3) a second phytoplankton bloom in autumn, followed by an increase in copepod density and a low macrofaunal abundance. In conclusion, pelagic and benthic communities in the coastal area of the Ligurian Sea mainly seem to be controlled bottom-up. Our results suggest that the quality of the particulate organic matter may play an important role in determining the temporal changes of both plankton and benthic assemblages, while the direct influence of other environmental features (such as sediment grain size) is relevant only for some macrobenthic taxa (e.g. crustaceans).     

Biochemical composition and early diagenesis of organic matter in coastal sediments of the NW Adriatic Sea influenced by riverine inputs

River inputs influence trophodynamic and biogeochemical processes of adjacent continental shelves. In order to provide new insights on the influence of continental inputs on the benthic trophic state and early diagenesis of sediment organic matter we collected surface sediments in the NW Adriatic Sea at three stations located at increasing distance from the Po River. Sediment samples were collected in four periods characterized by different river outflows and analysed for chloropigment content (chlorophyll-a and phaeopygments), protein, carbohydrate and lipid concentrations, prokaryote abundance and aminopeptidase activity. Sediments of the NW Adriatic Sea displayed high organic loads, tightly coupled with the outflow dynamics of the Po River. A major flooding event was responsible of an enhanced accumulation of organic material on the sea bottom. The resulting increased nutrient load in the sediment impaired organic matter degradation processes. The results of the present study suggest that the enhanced trophic state of marine coastal sediments subjected to riverine inputs are related not only to the increased nutrient inputs, but that they may be amplified by impaired degradation processes.     

Organic matter composition in sediments of the Oman Margin

The quantity and biochemical composition of organic matter (OM) in the sediments underlying and below oxygen minimum zone (OMZ) in the Arabian Sea were studied to provide information on the diagenetic processes of organic carbon under different environmental conditions. Concentrations of total organic carbon (TOC), total nitrogen (TN) and total hydrolysable amino acids (THAA) were significantly higher in sediments within rather than below OMZ, while those of total carbohydrates (TCHO) were slightly lower in the latter, suggesting the presence of a larger supply of labile compounds into the sea bottom at the shallower site (i.e. within OMZ). Hydrolysable amino acid and carbohydrate contribution to TOC were even lower (about 10% in surficial sediments) than the values obtained from the abyssal oligotrophic North Pacific, suggesting that OM food availability in the Oman Margin sediments within OMZ was lower than that observed at abyssal depths. The presence of the highest THAA and TCHO concentrations in the top 40?mm of the sediment core at both sites reflected the presence of bioturbation processes. In contrast with the general view of the deep sea as a stable and constant system, below OMZ in the Arabian Sea sediments, some differences were observed in the two investigated cores indicating the presence of a certain spatial variability in OM content and diagenesis.     

Organic matter composition in sediments of the Oman Margin

The quantity and biochemical composition of organic matter (OM) in the sediments underlying and below oxygen minimum zone (OMZ) in the Arabian Sea were studied to provide information on the diagenetic processes of organic carbon under different environmental conditions. Concentrations of total organic carbon (TOC), total nitrogen (TN) and total hydrolysable amino acids (THAA) were significantly higher in sediments within rather than below OMZ, while those of total carbohydrates (TCHO) were slightly lower in the latter, suggesting the presence of a larger supply of labile compounds into the sea bottom at the shallower site (i.e. within OMZ). Hydrolysable amino acid and carbohydrate contribution to TOC were even lower (about 10% in surficial sediments) than the values obtained from the abyssal oligotrophic North Pacific, suggesting that OM food availability in the Oman Margin sediments within OMZ was lower than that observed at abyssal depths. The presence of the highest THAA and TCHO concentrations in the top 40 mm of the sediment core at both sites reflected the presence of bioturbation processes. In contrast with the general view of the deep sea as a stable and constant system, below OMZ in the Arabian Sea sediments, some differences were observed in the two investigated cores indicating the presence of a certain spatial variability in OM content and diagenesis.     

Effect of temperature on biogeochemistry of marine organic-enriched systems: implications in a global warming scenario

Coastal biogeochemical cycles are expected to be affected by global warming. By means of a mesocosm experiment, the effect of increased water temperature on the biogeochemical cycles of coastal sediments affected by organic-matter enrichment was tested, focusing on the carbon, sulfur, and iron cycles. Nereis diversicolor was used as a model species to simulate macrofaunal bioirrigation activity in natural sediments. Although bioirrigation rates of N. diversicolor were not temperature dependent, temperature did have a major effect on the sediment metabolism. Under organic-enrichment conditions, the increase in sediment metabolism was greater than expected and occurred through the enhancement of anaerobic metabolic pathway rates, mainly sulfate reduction. There was a twofold increase in sediment metabolism and the accumulation of reduced sulfur. The increase in the benthic metabolism was maintained by the supply of electron acceptors through bioirrigation and as a result of the availability of iron in the sediment. As long as the sediment buffering capacity toward sulfides is not surpassed, an increase in temperature might promote the recovery of organic-enriched sediments by decreasing the time for mineralization of excess organic matter.     

对硝基苯酚在沉积物上的吸附特性1

研究了沉积物吸附对硝基苯酚的作用机理及影响因素.结果表明,对硝基苯酚在沉积物上的吸附由其中的有机质含量和结构决定,体系的pH值对对硝基苯酚的吸附有较大的影响,其主要原因是体系的pH值控制着沉积物中有机质的溶出量.在实验浓度范围内,分配作用在沉积物吸附对硝基苯酚中占主导地位.