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.     

Particulate Organic Matter Composition in A Semi-Enclosed Marine System

Spatial and temporal changes in the biochemical composition of particulate organic matter in a semi-enclosed marine system (Stagnone di Marsala, Mediterranean Sea) were studied, on a monthly basis, from January to December 1994, in order to assess nutritional value of suspended particles for benthic suspension feeders. According to previous findings, the study site displayed a strong oligotrophy. Chlorophyll-a accounted for a very low fraction of the total suspended matter pool (0.1%), whereas at least 75% of POC was of detrital/heterotrophic origin. POC: PON ratio values indicate that bacterioplankton biomass accounted for a significant fraction of the total POC pool, displaying values comparable to those of the phytoplankton biomass (phytoplankton to bacterial biomass ratio was about (1). Temporal and spatial changes in the biochemical composition of particulate organic matter were rather limited and related to its sources, the main of which is represented by detrital particles released by the Posidonia oceanica (L.) beds. the comparison between our results and those encountered in other coastal lagoons indicates that the low abundance of suspension-feeding organisms observed in the study area is related to the “quality depression” of particles due to the dilution of high quality compounds (i.e., biopolymeric carbon) in a largely inorganic matrix. This result leads us to conclude that, to reach the same amount of high quality particulate food, a suspension feeder mollusc in the Marsala lagoon would need to filter a sea water volume around 3 times higher than in other Mediterranean coastal lagoons.     

Changes in the Biochemical Composition of Tetraselmis Suecia and Isochrysis Galbana During Growth and Decay

The variations of the biochemical composition of Tetraselmis suecica and Isochrysis galbana during growth and decay were determined. the content of chlorophyll a (Chl-a) of the cultures, as expected, slowly degraded into phaeopigments during decay, confirming that chlorophyll measurements do not always provide an accurate estimate of phytoplanktonic biomass and, consequently, may fail if used to measure the food availability of particulate matter for consumers. Measurements of total amounts of proteins, carbohydrates and lipids, related to the nutritional value of particles in terms of caloric content, are shown to provide information on the readily available food for consumers, particularly during the blooms. the protein/carbohydrate, C/N and POC/Chl-a ratios were used to evaluate the differences between these two species during the growth and the decomposition processes. A comparison between experimental and field conditions was undertaken to implement our understanding of the growth and degradation processes of particulate organic matter of phytoplanktonic origin in the sea and its role on natural systems, during and after phytoplankton blooms.     

Particulate Organic Matter and Heavy Metals in Ligurian Open Sea

Seven stations were sampled along a transect between Imperia and Cape Corso during the C.N.R. “Bannock” survey (8-10 October 1984).

The TSM, PC, PN, and chlorophyll-a concentrations are low and decrease with depth. The AOU values suggest that below 400 m the organic matter is completely inert to further oxidation. Heavy metals concentrations show marked differences in the surface water layer between coastal and offshore stations. The chemical characteristics of total suspended matter are more conservative as an index of waters of coastal origin than the quantity of particulate matter itself or metal concentrations. The Fe: Cd ratio in particles is of interest as an indicator of coastal waters.     

A detailed analysis of carbohydrates in marine particulate matter

Detailed profiles of organic carbon, organic nitrogen, carbohydrate and proteinous amino acid have been determined in particulate matter from various depths at a station in Sagami Nada off eastern Honshu, Japan. The profiles suggest that carbohydrate is decayed from particulate matter more rapidly than proteinous amino acid in the euphotic zone, while the latter is removed to a higher extent than the former in deep waters,which results in the increase of the C/N values of particulate matter in the depths. All of the particulate matter gave D-galactose, D-glucose, D-mannose, D-xylose and D-glucuronic acid upon acid hydrolysis. Vertical change of the monosaccharide composition of this particulate matter indicates that only D-glucose and its polymers are preferentially removed from it during its descent. Carbohydrates from the particulate matter of 20 m depth were fractionated into water-soluble and insoluble carbohydrates. Detailed analysis of these fractions indicates that 1,3-glucan and its related low molecular weight carbohydrates from the water-soluble fraction decay between 50 and 300 m depth, to leave waterinsoluble carbohydrates, immune to biological attack during the course of further sinking. On the basis of these facts, the effects of the biochemical nature of these carbohydrates on the vertical change of the particulate carbohydrate distribution are discussed.     

Origin and Distribution of Suspended Organic Matter As Inferred From Carbon Isotope Composition in A Mediterranean Semi-Enclosed Marine System

The origin and distribution of suspended organic matter, the trophic features and the stable carbon isotopic composition of particulate organic carbon (POC) were studied monthly in a Western Mediterranean semi-enclosed basin. Sampling stations were selected as a function of wind-exposure and the degree of vegetation cover and then compared with an adjacent unvegetated site. the predominant vegetation was seagrass (Posidonia oceanica and Cymodocea nodosa) and Caulerpa prolifera. Water samples were analyzed for total suspended matter (inorganic and organic fractions), photosynthetic pigments (chlorophyll-a and phaeopigments), dissolved organic carbon, particulate organic carbon and their isotopic composition. Temperature and salinity were also measured at the same sampling sites within range of Mediterranean limits. the suspended organic matter concentration was 1.77 ± 1.55 mg l^-1; the chlorophyll-a concentration was low (0.35 ± 0.24 μg l^-1); the disolved organic carbon concentration was 2,140 ± 2,010 μg l^-1; the particulate organic carbon concentration was 212 ± 106 μg l^-1 and the isotopic composition was 18.77 ± 2.51%_°. There were significant temporal differences except for phaeopigments, POC and its POC isotopic composition, and there were no spatial differences other than for δ¹³C. This picture highlighted a general seasonal trend and trophical features similar to adjacent sea.

Spatial differences in δ¹³C showed that the source of suspended organic matter was different between stations as that between sources and wind-hydrodynamic constraints. In     

Total and “Selective” Extraction Methods for Trace Metals in Marine Sediment Reference Samples (Mess-1, NBS 1646)

Trace metals (Cd, Cr, Cu, Mn, Pb and Zn) and Fe were analyzed in two sediment reference samples (NBS 1646, MESS-1) with AAS fitted with graphite furnace, using five different (three total and two selective) extraction procedures.

The comparison of the total digestion methods (hot strong acids) with reference materials gave good results for almost all of them, and the “teflon bomb” was preferred for its rapidity and ease of operation. Some non-referenced data on total and organic carbon, total nitrogen and minerals are given. The two selective leaching extractions (nitric and cold hydrochloric) were also compared with reference values.

The use of certified reference materials (CRM) for environmental chemistry is recommended, together with the determination of organic matter and fundamental mineralogical composition.     

Organic and Inorganic Characterization of Marine Colloids

Isolates of the organic matter in the particulate, colloidal and dissolved states were obtained by tangential flow ultrafiltration through 0.40 μm polycarbonate and ∼ 1 nm (1000 NMWL*) regenerated cellulose membranes and by solid-liquid reverse phase extraction techniques. the material was analyzed qualitatively by mass spectroscopy, proton nuclear magnetic resonance, Fourier transform infrared spectroscopy, high performance liquid chromatography with fluorescence detection of labelled primary amines and amino acids, and inductively coupled plasma/mass spectroscopy. All three states are characterized by similar organic chemistries. the marine colloidal state in coastal waters off the Californian coast contains primarily carbohydrates, fatty acids, minor amounts of proteinaceous compounds and electropositive elements including aluminium and iron. Aromatic molecules and olefinic functional groups are in low concentration. the colloidal state differs qualitatively from the particulate and dissolved states. Yet all three could be derived, with degradation, from algal or macroalgal surface components.     

Sediment organic matter and meiofauna community response to long-term fish-farm impact in the Ligurian Sea (Western Mediterranean)

Quantitative and qualitative changes in meiofauna community structure were investigated to assess the impact of a fish farm, which was operating continuously for 15 years (La Spezia Gulf, W Mediterranean). Sediment samples were collected in June, July, September, October 2000 and February 2001 for the analysis of phytopigments (chlorophyll-a and phaeopigments), the biochemical composition of organic matter (proteins, carbohydrates and lipids) and related to meiofaunal parameters.

Sediment organic matter reached extremely high concentrations beneath the fish cages when compared to the control. Particularly lipids, carbohydrates and chlorophyll-a were significantly higher in fish-farm sediments. On a long-term basis meiofauna displayed adaptations in sediments beneath the cages resulting in an increase of density. Organic impact on meiofaunal community structure was evident in terms of an increase of the nematodes to copepods (Ne/Co) and nauplius to copepods (Na/Co) ratios in fish farm sediments. Cumaceans and kinorhynchs were encountered in control sediments, but disappeared in fish-farm samples. These data suggest that meiofauna is a sensitive tool for evaluating the effects of organic enrichment in fish farm impacted areas.     

Variability of biochemical composition and size distributions of seston in the euphotic zone of the Bay of Biscay: implications for microplankton trophic structure

Particle-size distributions and several biochemical components of seston were studied from October 1986 to December 1987 in surface waters of the Bay of Biscay. Variance partitioning of hydrographic and seston parameters indicates that, although seasonal variability related to the thermal cycle was important, most of the differences in seston concentration were caused by spatial components of variance. Changes in the vertical structure of the water column appeared to be the principal source of variability. Phytoplankton blooms in spring and fall and the effects of the coastal upwelling and the thermocline during summer were traced using seston concentration and biochemical composition. The different seston measurements were scaled according to their correlations with total concentration and particle size. Photosynthetic pigment biomass was related to both large and small partieles. However, concentrations of particulate proteins, lipids and carbohydrates were more related to small particles. Although not specifically analysed, low concentrations of inorganic particles and detritus can be expected in the study area, thus most of the particulate organic matter was associated with small, non-pigmented organisms, particularly in surface waters during the period of thermal stratification. The variability in concentrations of total seston and particulate organic matter was mainly due to variations in particulate proteins, lipids and carbohydrates; pigment concentrations were of secondary importance. Expressed as the ratio chlorophyll a: particulate protein-nitrogen, phytoplankton constituted an important fraction of the microplankton biomass only during spring blooms, when it averaged 75% of the particulate protein-nitrogen. In constrast, <30% of protein-nitrogen was related to chlorophyll a during the summer. These results suggest that an oligotrophic structure based on the microbial-loop prevails in microplankton assemblages of surface waters for most of the year.     

Studies on water-extractable carbohydrates of the particulate matter from the northwest Pacific Ocean

We have studied carbohydrate metabolism in the marine environment by means of the isolation and characterization of carbohydrates of particulate matter from various depths. The present work is especially concerned with water-extractable carbohydrates of the particulate samples from 11 stations of the northwest Pacific Ocean. Water-extractable carbohydrate content in the particulate matter of the surface waters was measured; it ranged from 19.2 to 36.0% of the total particulate carbohydrate. These values decreased, ranging from 6.5 to 15.5% at 50 m depth. Such rapid vertical change of the concentration of the carbohydrate indicated that it was lost from the particulate matter much faster than the other particulate organic constituents such as protein and water-insoluble carbohydrate. This process left water-insoluble carbohydrate in the deep-water particulate matter as previously observed in the Sagami Nada, off eastern Honshu, Japan. Upon acid hydrolysis, water-extractable carbohydrate revealed glucose ranging from 67.0 to 84.2%. Rhamnose, fucose, ribose, arabinose, xylose, mannose and galactose were also identified as components of the carbohydrate fraction. Carbohydrate studies on the laboratory cultured diatoms clearly indicated that the water-extractable carbohydrates were food reserve materials of the diatoms. In view of this fact, the metabolism of the water-extractable carbohydrate in the euphotic zone and its underlying layer is discussed.     

Seasonal Fluctuations in the Nutritional Value of Particulate Organic Matter in A Lagoon

Water samples were collected on a fortnightly basis in the lagoon of S. Gilla (Sardinia, Mediterranean Sea) in order to study seasonal nutritional fluctuations of particulate organic matter. the lagoon is characterized by high quantities of suspended matter throughout the year. Thermohaline conditions had no effect on particulate matter quantity and composition, but the quantity as well as quality of suspended particles was drastically affected by the wind, the major effecter of sediment resuspension. As a result of sediment resuspension, seston was always richer in inorganic fraction. However, throughout the year of investigation, most particulate organic carbon was quite appealing for filter feeding communities, although the best POM quality was available during phytoplankton blooming. the phytoplankton pool of suspended matter was just a small fraction of the bulk, accounting for only 13% on average of particulate organic carbon. in terms of energy available in the seston, the highest amount was stored in organic matter heterotrophic fraction, whilst the smallest was to be found in living phytoplankton.     

Scope for growth of Mytilus?galloprovincialis (Lmk., 1819) in oligotrophic coastal waters (Southern Tyrrhenian Sea, Italy)

The ‘scope for growth’ (SFG) tool was used to study the growth performance of cultivated populations of Mytilus galloprovincialis (Lmk., 1819) in an oligotrophic area of the Southern Mediterranean Sea. The study was carried out between 1993 and 1996 by using data from four seasonal oceanographic cruises and from growth experiments. Water samples were collected and analysed for total suspended matter (TSM), particulate organic carbon (POC) and nitrogen (PON), particulate lipids, proteins and carbohydrates and chloropigments. The sum of the carbon equivalents of carbohydrates, proteins and lipids is indicated as the total biopolymeric particulate organic carbon (BPC) and was converted into a unit of energy in order to calculate the SFG of a theoretical mussel of 5 cm length. In order to test the performance of mussel growth at two depths (5 and 15 m water depth), mussel body size [as ash free dry weight (AFDW)] and the actual concentrations of BPC were used to calculate the monthly SFG using the physiological energetic relationships suggested in the current literature. Data from the field cruises led us to characterise the study site as ultra-oligotrophic (annual average of chloropigment concentration approximately 0.5 μg L⁻¹). SFG calculations allowed us to identify a site where mussels grown successively were found to reach a commercial size in approximately 12 months. The good agreement obtained between energetic response and subsequent production response suggests that the available energy from particulate food could be fully available for organic production for maintaining “proportionate” growth trajectories, even in a ultra-oligotrophic system.     

Sources and biochemical composition of suspended particulate material in a submarine cave with sulphur water springs

Sources, biochemical composition and nutritional value of suspended particulate material were investigated from February 1994 to February 1995 in a submarine cave (Grotta Azzurra, Capo Palinuro, southwestern Italy) with hot sulphur springs and associated mats of chemolithoautotrophic bacteria in its innermost dark part (Snow Hall). Concentrations of total suspended material (TSM), particulate inorganic material (PIM), organic carbon (POC), organic nitrogen (PON), chlorophyll a (Chl a), phaeopigments (Phaeo), carbohydrates (TCH), proteins (TPR) and lipids (TLI) were measured at four stations along an outside–inside transect, in order to address whether the quantity and quality of suspended particles varied over time with increasing distance from the entrance of the cave and estimate the relative contribution of chemosynthesis versus photosynthesis in supplying POC to cave heterotrophs. The abundance and biochemical composition of suspended material available to filter-feeders varied over time, but no significant quantitative or qualitative differences were detected along the outside–inside transect. Concentrations of TSM and of its different compounds (PIM, POC, PON, Phaeo, TCH, TPR, TLI) were homogeneous among the four stations or fluctuated without any consistent trend, with no apparent ageing and degradation of organic material in the innermost dark station. Conversely, concentrations of Chl a significantly decreased from outside to inside the cave at all sampling periods. It is suggested that suspended POC in Grotta Azzurra consists of mixed assemblages of particles coming from advection of photosynthetic material from the open sea and local inputs of carbon by sulphur-oxidizing bacteria. Based on POC/Chl a ratios, the relative contributions of chemosynthesis versus photosynthesis in supplying POC to benthic heterotrophs was estimated to be 31 and 69%, respectively. Received: 9 December 1996 / Accepted: 14 January 1997     

Sediment organic matter and meiofauna community response to long-term fish-farm impact in the Ligurian Sea (Western Mediterranean)

Quantitative and qualitative changes in meiofauna community structure were investigated to assess the impact of a fish farm, which was operating continuously for 15 years (La Spezia Gulf, W Mediterranean). Sediment samples were collected in June, July, September, October 2000 and February 2001 for the analysis of phytopigments (chlorophyll-a and phaeopigments), the biochemical composition of organic matter (proteins, carbohydrates and lipids) and related to meiofaunal parameters.

Sediment organic matter reached extremely high concentrations beneath the fish cages when compared to the control. Particularly lipids, carbohydrates and chlorophyll-a were significantly higher in fish-farm sediments. On a long-term basis meiofauna displayed adaptations in sediments beneath the cages resulting in an increase of density. Organic impact on meiofaunal community structure was evident in terms of an increase of the nematodes to copepods (Ne/Co) and nauplius to copepods (Na/Co) ratios in fish farm sediments. Cumaceans and kinorhynchs were encountered in control sediments, but disappeared in fish-farm samples. These data suggest that meiofauna is a sensitive tool for evaluating the effects of organic enrichment in fish farm impacted areas.     

Biodiversity and trophic structure of nematode assemblages in seagrass systems: evidence for a coupling with changes in food availability

Nematode assemblage composition, trophic structure and biodiversity were followed over an annual cycle in a Posidonia oceanica bed of the NW Mediterranean to test the response to temporally changing food availability (sediment organic matter, bacterial and microphytobenthic biomass). The sediment-water interface of the seagrass meadow was characterised by high particulate organic matter concentrations. Also, seagrass sediments had high organic matter content, chloropigment concentrations and bacterial biomass. All organic matter components (i.e. carbohydrates, proteins and lipids) changed temporally, with higher concentrations in winter and spring and lower concentrations in summer; however, overall, large amounts of organic compounds were potentially available for (seagrass) benthic consumers throughout the year. Nematode assemblage in the P. oceanica bed maintained a high genus number (88 genera), and a trophic structure tightly coupled with the composition of the potential food sources. In agreement with the relevance of microphytobenthos, epigrowth-feeder nematodes (2A) represented the dominant trophic guild. The biomass of predator nematodes (2B) was significantly correlated with the biomass of other nematodes. Non-selective deposit feeders (1B) displayed a close relationship with the concentration of proteins, carbohydrates and biopolymeric carbon (i.e. labile organic detritus). Conversely to what was expected, epigrowth feeders were not correlated with chloropigment concentrations, but showed a significant relationship with the number of dividing bacteria, possibly suggesting a trophic plasticity of the 2A feeding guild. The coupling between temporal changes in food sources and temporal variability of the nematode trophic structure was highlighted by a CANOCO analysis, which allowed us to identify and associate, at each sampling time, nematode genera and their feeding habits, with specific environmental variables and food indicators. Furthermore, species diversity (H') and evenness (J) calculated on nematodes identified to the genus level displayed evident temporal changes, also reflected by the index of trophic diversity (ITD). Both structural and functional diversity were coupled with high concentration and highly heterogeneous composition of the food sources (including organic detritus, microphytobenthic algae and bacteria) potentially available to nematodes. These data suggest that temporal changes in quantity and quality of food sources do not only influence nematode dynamics and trophic composition, but also influence nematode structural and functional diversity.     

Decomposition of faecal matter and somatic tissue of Mytilus galloprovincialis: changes in organic matter composition and microbial succession

Variations in the biochemical composition of pseudo-faeces and faeces egested by Mytilus galloprovincialis (Lamarck) and in detritus derived from the somatic tissue of mussels during the decomposition process were investigated by means of two intensive experiments. During the degradation process, the biochemical composition of pseudo-faeces and faeces showed a clear increase in protein content related to the microbial colonization. Changes also occurred in the biochemical composition of particulate organic matter (POM) in the surrounding water due to faecal matter decomposition, heterotrophic utilization and conversion of particulate carbohydrates and proteins to the dissolved pool. The study of production and heterotrophic utilization of the POM derived from the somatic tissue of M. galloprovincialis collected in the Gulf of Tigullio, Italy in 1990 indicates that this kind of material is rapidly decomposable and largely available for benthic consumers. Bacteria utilized selectively the different compounds, and proteins proved to be the most suitable substrate for bacterial growth. The input of organic detritus into the experimental system resulted in an enhancement of bacterial activity and consequently of the RNA/DNA ratio. Bacterial DNA accounted on average for 17% of particulate DNA. During decomposition processes, nutrient release was about ten times higher than the value calculated from individual excretion rates, indicating that mussel beds may be important sites for nutrient regeneration. Carbon conversion efficiency for bacteria growing on faecal matter was, on average, 17.2%. The potential importance of faecal output and bacterial production as a carbon resource for benthic communities near mussel culture areas is discussed.     

Biochemical composition of sedimentary organic matter in the coral reefs of Lakshadweep Archipelago,Indian Ocean

Surface sediments were collected from the shore and lagoons of Kavaratti, Kadamat Agatti and Pitti islands of Lakshadweep Archipelago during May 2015 and analysed for biochemical composition and quality of organic matter. The biochemical composition of sedimentary organic matter from the entire study area was characterised by the dominance of carbohydrates (CHO) followed by proteins (PRT) and finally lipids (LPD). PRT:CHO ratios were less than 1 and indicated the presence of aged organic matter in the islands. The poor nutritional quality of sediments to support benthic fauna was evident from the values of LPD:CHO ratios. The refractory nature of sediments and less availability of food to benthic source was supported by BPC:TOC ratios. Based on estimated ratios and biopolymeric carbon values, the trophic status of the study area was categorised as oligotrophic.     

Organic and Inorganic Characterization of Marine Colloids

Abstract

Isolates of the organic matter in the particulate, colloidal and dissolved states were obtained by tangential flow ultrafiltration through 0.40 μm polycarbonate and ～ 1 nm (1000 NMWL?) regenerated cellulose membranes and by solid-liquid reverse phase extraction techniques. the material was analyzed qualitatively by mass spectroscopy, proton nuclear magnetic resonance, Fourier transform infrared spectroscopy, high performance liquid chromatography with fluorescence detection of labelled primary amines and amino acids, and inductively coupled plasma/mass spectroscopy. All three states are characterized by similar organic chemistries. the marine colloidal state in coastal waters off the Californian coast contains primarily carbohydrates, fatty acids, minor amounts of proteinaceous compounds and electropositive elements including aluminium and iron. Aromatic molecules and olefinic functional groups are in low concentration. the colloidal state differs qualitatively from the particulate and dissolved states. Yet all three could be derived, with degradation, from algal or macroalgal surface components.     

Spatial and Temporal Changes of Suspended Matter in Relation to Wind and Vegetation Cover in A Mediterranean Shallow Coastal Environment

Seasonal and spatial changes in seston, (POC), particulate organic carbon, (PON) particulate organic nitrogen and chlorophyll-a concentrations were studied on a monthly basis in a Mediterranean shallow coastal area (Stagnone di Marsala, Western Sicily) in order to gather information on factors controlling particulate organic matter distribution and composition. Seston concentration and composition were connected to the main physicochemical and biological driving factors, such as temperature, salinity, dissolved oxygen, wind-speed and biomass of submerged vegetation. the Stagnone di Marsala is characterized by high temperatures with strong seasonality (range: 11-28°C), while values ranged from 33 to 45 salinity. Total suspended organic matter concentrations (by ignition loss) ranged from 2 mg l^-1 (in summer) to 12mgl^-1 (in winter) and chlorophyll-a concentrations from 0.02 to 2 μgl^-1. Despite a low POC/PON ratios (ranging from 5 to 11), the ratio of POC to chlorophyll (CHL-a) displayed very high values (annual average of 647). the data reported in this study, highlighting the oligotrophy of the Stagnone di Marsala area, indicate that the trophic state of the basin was controlled by different degrees of wind exposure (mean monthly wind velocity at exposed sites ranged between 4.2 and 6.7 m s^-1) and by gradients in vegetation cover. These two factors induced clear changes in the concentration and composition of the suspended particles, but played a different role in exposed and sheltered areas. Exposed areas with limited vegetation were characterized by large resuspension processes and wide temperature and salinity fluctuations caused by wind induced turbulence. in these areas, autotrophic biomass (as chlorophyll-a), due to phytoplankton and/or re-suspended microphytobenthos, appeared to play an important