Patterns of micro- and meiofaunal abundance in marine sediments,measured with the adenosine triphosphate assay

Adenosine triphosphate (ATP) measurements are used to determine vertical and seasonal distributions of microorganisms and meiofauna in sediments from a 14 m-deep mud bottom in central Long Island Sound on 12 sampling dates from April 1975 to October 1976. Below the topmost 1 cm of sediment, ATP measurements can be useful in estimating and comparing standing stocks of microorganisms and meiofauna. In the top 1 cm, however, large quantities of newly settled bivalves (Yoldia limatula, Nucula annulata, and Mulinia lateralis) and juvenile polychaetes (Owenia fusiformis) in summer and fall months account for total ATP concentrations. The ATP content of individual meiofauna ranges from 1.97 ng individual copepod nauplius^-1 to 190.7±60 ng individual M. lateralis ^-1. In general, the total ATP content of individual polychaetes and bivalves is much higher than that of individuals of other groups. However, on a mg ATP per g wet or dry tissue basis, the ATP content of micro- and meiofaunal taxa are not significantly different. In addition to providing a means for comparing micro- and meiofaunal standing stocks, ATP measurements permit examination of the relative contribution of different meiofauna to the total living biomass of meiofauna in sediments. Total sediment ATP concentrations are greatest in the top 1 cm at all seasons, and decrease with increasing depth in the sediment. Annual concentrations in the topmost centimeter average 4.22 g g dry sediment^-1 and range seasonally from 1.09 to 7.64 g g dry sediment^-1. At a depth of 10 cm, values average 0.16 and range from 0.019 to 0.35 g g dry sediment^-1. High ATP concentrations in surface sediment reflect high concentrations of microorganisms and meiofauna at the sediment-water interface. The top 2 cm of sediment contain 71% of all meiofauna, with 41% occurring in the topmost cm. In general, densities are lowest in the winter and highest during the spring and summer, averaging 490 individuals 10 cm^-2, and varying from 87 to 1366 individuals 10 cm^-2. Because of wide variation in recruitment patterns of the benthos in Long Island Sound, the extremes of the range in meiofaunal densities can be observed in the same month in two different years. In order to monitor and compare standing stocks of organisms less than 1 mm in size in sediments, the ATP assay can save hours of processing time compared with alternate methods such as direct counts.     

Experimental variation of near-bottom current speeds and its effects on depth distribution of sand-living meiofauna

Meiofauna are eroded from sediments and transported in the water column. To examine how meiofauna respond to erosive currents, in situ current speeds were altered on an intertidal sandflat located in Bogue Sound, North Carolina, USA in July 1982. Replicates of two different weir designs were erected. One design increased measured current speed and the other, a control, did not. The depth distributions of meiofauna in each treatment and from undisturbed sediment were compared. There were few differences among treatments in depth distribution of the most abundant taxa: nematodes, turbellarians, and ciliates (>37-m-long). However, a significant amount of sediment erosion (a mean of almost 0.5 cm in 2 h) occured in the increased flow treatment. Meiofauna responded to the loss of sediment with no discernable differences in depth distribution relative to the controls. There was some evidence that nematodes were less abundant and ciliates more abundant in the top 0.5 cm of the increased flow treatment. This may indicate that nematodes respond to fast surface current speeds by moving deeper into the sediments, while ciliates manage to remain near the surface even as sediment erodes. The entrainment of meiofauna may be passive but meiofauna clearly exert considerable behavioral influence over their susceptibility to entrainment.     

Determination of average individual dry weights and ash-free dry weights in different sieve fractions of marine meiofauna

Individual dry weights and ash-free dry weights for different meiofauna taxa from two soft bottom localities (31 and 117 m depth) in Gullmar Fjord on the Swedish west coast were determined. Samples from August 1980 and February 1981 of about 100 to 600 formalin-preserved specimens of each of the different taxa from each of four sieve fractions were weighed on a microbalance. Individual dry weights ranging from 0.12 g (nematodes in the 0.040-mm fraction) to 172 g (polythalamous, arenaceous Foraminifera in the 0.500-mm fraction) and ash-free dry weights ranging from 0.10 g (nematodes in the 0.040-mm fraction) to 37 g (polychaetes in the 0.500-mm fraction) were obtained. Great similarities were noted between corresponding values from the two stations as well as between summer and winter values. The percentage of organic matter, determined by ashing, in the body weight differed greatly among taxa. Some standard individual dry weight and ash-free dry weight values, to be used for re-calculation of abundance values into biomass, are presented.     

Benthic response to sedimentation events during autumn to spring at a shallow water station in the Western Kiel Bight

Seasonal variations in bacterial populations (total number, biomass, biomass-spectrum, number of dividing cells) as well as in concentrations and decomposition rates of particulate organic material were followed in a sandy mud sediment of the Western Kiel Bight (Baltic Sea; FRG). The strong seasonal variations observed could be traced back to the effect of certain ecological situations and events in the sediment from which the input of the phytoplankton blooms in autumn and spring, respectively, the accumulation of organic material during winter, and the spring development of the benthic fauna turned out to be the most important. Bacterial carbon net production following the breakdown of the phytoplankton blooms ranged between 9 g (autumn) and 16 g (spring) per g of dry weight sediment per day. The consequences of shifts in the size composition of the bacterial populations as well as the importance of the measurement of enzymatic decomposition rates of particulate organic material in sediments are demonstrated and discussed in relation to the events mentioned above.Publication No. 420 of the Joint Research Program at Kiel University (Sonderforschungsbereich 95 der Deutschen Forschungsgemeinschaft)     

Rates of seasonal sediment reworking in Axiothella rubrocincta (Polychaeta: Maldanidae)

Sediment reworking rates of Axiothella rubrocincta (Johnson, 1901) (Polychaeta: Maldanidae) were measured in situ in Tomales Bay, California (USA), from August, 1969 through July, 1970. On the average, each adult worm (approximate fresh weight=1 g) reworks about 5 g dry sediment d^-1 at a mean temperature of 13.4°C and a mean salinity of 31.8.Reworking rates are positively correlated with temperature and salinity, and negatively with sediment organic carbon, sedimentation rates and grain size. An inverse correlation exists between sediment reworking rates (g dry sediment g^-1 wet wt of worm d^-1) and g wet weight of worm. Sediment parameters deseribing unworked sediments are not significantly different from those for fecal sediments. Although these data suggest this species to be a non-selective deposit-feeder, it is more likely that it is faculatively selective.     

Quantitative estimates of the meiofauna from the deep sea off North Carolina,USA

In 1974 the authors collected, sorted and enumerated meiofauna from 400, 800 and 4000 m off North Carolina, USA. Samples were replicated respectively with 4 boxcores and 21 subsamples, 2 boxcores and 7 subsamples and 2 boxcores and 8 subsamples. Total meiofaunal numbers were highest in fine silt sediment at 800 m ( =891.9 10 cm_–2) and lowest in very fine silt at 4000 m ( =73.5 10 cm_–2). Fine sand at 400 m yielded a mean of 442.4 10 cm^–2. At all depths, most fauna were located in the upper 3 cm of sediment ( depth distribution=2.2 cm), and typically only nematodes and foraminiferans were found below 4 cm. Total community abundances significantly differed with depth; however, there were no differences among replicate boxcores at particular depths. Since most (85.7%) of the variance was associated with subsamples from a boxcore, it appears that meiofauna densities are homomeneous within large areas at particular depths, and that patchiness is a smallscale phenomenon at the level of the 10 cm² subsampler. Comparisons of sorting efficiencies of live and preserved samples indicated that to accurately enumerate formainiferans, samples must first be fixed and stained, while turbellarians and oligochaetes must be sorted live.Contribution No. 151 from the Belle W. Baruch Institute for Marine Biology and Coastal Research. This research was supported by the Oceanography Section, National Science Foundation, NSF Grant GA-42792, and GC-00005 to Duke University for R.V. Eastward operations.     

Seasonal changes of benthic bacteria in a seagrass bed (Posidonia oceanica) of the Ligurian Sea in relation to origin,composition and fate of the sediment organic matter

Variations in number and biomass of benthic bacteria were examined in the surface sediments of a Mediterranean seagrass bed [Posidonia oceanica (L.) Delile] in the Gulf of Marconi (northwestern Mediterranean Sea) from 1990 to 1991. The annual dynamics of benthic bacterial density and biomass were compared to changes in elemental (organic C and total N) and biochemical (lipids, proteins, carbohydrates) composition of sediment organic matter, as well as to microphytobenthic biomass, dissolved inorganic nutrients and ATP. Bacterial densities exhibited marked seasonal variations (5.12 to 322.7x10⁸ cells g^-1 sediment dry wt) with highest values in late spring. Bacterial standing stocks (15.8 to 882.33 g C g^-1 of sediment dry wt) were high. Bacterial biomass did not correlate with organic C, total N or to specific biochemical components, but correlated significantly with chlorophyll a, ATP and porewater phosphate concentrations. There is evidence that benthic bacteria were responding to variations of algal biomass. Bacterial biomass accounted, on average, for 30% of total living carbon (calculated on the basis of the ATP concentrations) and 8.4% of total organic carbon.     

Nitrogenous excretion by meiofauna from coral reef sediments: Mecor 5

Nitrogenous excretion rates of meiofauna from coral sediments of the Great Barrier Reef, Australia were studied at winter temperatures. Data on single taxa fitted a geometric regression: Log N excretion=-0.206+0.865 log x where log x=dry weight in g. Abundance and biomass data on meiofaunal taxa were obtained from Lizard Island, Orpheus Island and Davies Reef. Using excretion rates and biomass data, meiofauna were calculated as contributing 10.18, 12.32, and 1.60 mg N m^-2 h^-1 at the above areas respectively.Contribution No. 5 from the workshop Microbial Ecology of coral Reefs (MECOR), sponsored by the Australian Institute of Marine Sciences and the US National Science Foundation, July–August 1984     

ATP concentration and adenylate energy charge of tropical zooplankton from waters inside the great barrier reef

The adenylate energy charge ratio (EC) of 8 species of zooplankton ranged from 0.70 to 0.89. Eucalanus subcrassus had significantly (P<0.05) lower EC when captured during a 30-min tow than during a 3-min tow, indicating stress at capture. The concentration of ATP in the 8 species ranged from 9.1 to 36.0 g/mg protein and from 0.17 to 15.7 g/mg dry weight. Excluding a low value for the ctenophore Pleurobrachia sp., the average concentration was 8.1 g ATP/mg dry weight, which is equivalent to an approximate C/ATP ratio of 50.Contribution from the Australian Institute of Marine Science, Australia, and the Institute of Marine Biology, University of Bergen, Norway     

Brood size and chemical composition of Pareuchaeta norvegica (Crustacea: Copepoda) in Loch Etive,Scotland

Brood size and chemical composition (carbon, nitrogen, and ash) of Pareuchaeta norvegica in Loch Etive, Scotland have been determined. From 23 to 37 eggs were found in each egg mass (mean=30 eggs). The eggs are oval; the mean size of the longer axis is 438 m. Variation in egg mass (as dry weight) is related to increase in dry weight of the maternal body. an increase of 1 mg in maternal body dry weight produces an increase of 0.03 mg dry weight in the egg mass. Carbon content increases with increasing body dry weight both in pre-spawning and spent females, but nitrogen content decreases with increasing body weight. Thus, the carbon:nitrogen ratio increases with increasing body dry weight. The carbon content of an egg mass ranges from 61.4 to 65.3%, and nitrogen content from 9.6 to 13.2% of the total dry weight. Some examples of partitioning of chemical components of the maternal body into the egg mass are also described.     

Effect of pentachlorophenol (PCP) on meiobenthic communities established in an experimental system

Aquaria containing clean sand received a continuous supply of seawater from Santa Rosa Sound, Florida, USA, mixed with known concentrations (7, 76 and 622 g l^-1) of pentachlorophenol (PCP). After 9 weeks, nematodes accounted for 87% of the total meiofauna. Nematode biomass and densities were greatest in aquaria exposed to 76 g PCP l^-1 and were least in aquaria exposed to 622 g PCP l^-1. Epistrate feeders were abundant in control aquaria and aquaria exposed to 7 and 76 g PCP l^-1, but not in aquaria exposed to 622 g PCP l^-1. Selective deposit feeders were not abundant in the control aquaria and aquaria exposed to 7 g PCP l^-1, but comprised 19% of the nematodes in aquaria exposed to 76 g PCP l^-1 and 61% in aquaria exposed to 622 g PCP l^-1.     

Elemental composition (C,N,H) in larval and crab-1 stages of Pagurus bernhardus (Decapoda,Paguridae) and Carcinus maenas (Decapoda,Portunidae)

Analyses of individual content of carbon (C), nitrogen (N), and hydrogen (H) were carried out for all larval stages of Pagurus bernhardus and Carcinus maenas, and for newly metamorphosed crabs. Maximum range in total larval development is 12.8 to 165.8 g C, 3.2 to 35.1 g N, and 1.9 to 24.9 g H in P. bernhardus and 3.1 to 43.2 g C, 0.7 to 10.1 g N, and 0.4 to 6.3 g H in C. maenas. From these data energy equivalents were calculated. Maximum range in total larval life is 0.43 to 6.38 J ind. ^-1 in P. bernhardus and 0.1 to 1.49 J ind. ^-1 in C. maenas. There is a 32.4% mean loss of energy in P. bernhardus megalopa development; this seems to describe the normal developmental pattern in this stage. Biomass was determined as fresh and dry weight respectively. Individual dry weight is about 3.6 to 5.6 times higher in P. bernhardus (44 to 340 g) than in C. maenas (12 to 93 g) larvae.Contribution to research project Experimentelle marine Ökosystemanalyse sponsored by Bundesministerium für Forschung und Technologie, Bonn (Grant No. MFU-0328/1)     

Benthic response to sedimentation of a spring phytoplankton bloom: Process and budget

The response of benthos to sedimentation of the spring phytoplankton bloom in the Kiel Bight (Western Baltic Sea) is described in terms of biomass (ATP) and activity (heat production and ETS-activity). Input of the bloom (11.5 g C m^-2) over a period from March 25 to April 19, 1980 to the sediment surface was in the form of cells and fresh phytodetritus as indicated by low C/N ratios (7) and high energy charge values (0.78). Benthic microbial activity was immediately stimulated by this input as heat production doubled and the activity of ETS tripled over winter values within 12 d in the absence of a significant increase in ambient temperature. A comparison of the two activity parameters suggests that anaerobic metabolism is more important during the winter (February and March) than after input of the bloom. Meiofauna was not able to take part in the first activity outburst. Benthic ATP-biomass (excluding macrofauna) doubled in late April due to microbial production, and doubled again in early May when meiofauna started reproductive activity. For macrofauna a general statement was not possible, although the sediment surface feeder Macoma baltica commenced a build up of glycogen and lipid resources immediately following bloom input whereas Nephtys ciliata, feeding on sediment and small macrofauna, showed a less pronounced and delayed effect from this input. An energy budget based on heat production measurements was calculated. A daily heat loss of the benthic community of 21.7 KJ m^-2 d^-1 (35.5 KJ m^-2 d^-1) was found, when a depth of 3 cm sediment (5 cm) was assumed. Heat production of macrofauna contributed less than 5% of this activity. The input of the bloom was burned within 21 (13) d. Preliminary estimations for an annual budget suggest that the vertical transport of particulate organic matter via sedimentation can only explain 25% (15%) of the benthic activity in the shallow water ecosystem of the Kiel Bight. This indicates the presence of other sources of organic carbon such as benthic primary production or other transport processes providing carbon to the sediments.Publication No. 384 of the Joint Research Program of Kiel University (Sonderforschungsbereich 95)     

Some biologically important low molecular weight organic acids in the sediments of Loch Eil

The isolation of biologically important low molecular weight organic acids from organically enriched sediments in Loch Eil, Scotland, was carried out by extraction of pore water with acidified ethyl acetate. High concentrations of acetic acid, up to 1.8 mg g^-1 dry weight of sediment were found at Station E-24. Propionate, butyrate, valerate, lactate and traces of succinate were also found. Succinate was present in significant amounts, 42.2 g g^-1 dry weight of sediment at Station E-70, which received a higher input of organic matter than E-24. Both propionate and succinate were absent from a control station in the Lynn of Lorne where the sediment was low in carbon compared with Loch Eil. In experimental tanks, acetate levels increased as the input of organic carbon (as cellulose) was increased up to a load level of 1.5 g m^-2 day^-1. Above this, acetate decreased and succinate appeared. Succinate was not detected in low-loaded tanks. Experiments with sieved mud showed a vertical distribution of the different acids with depth. Lactate and succinate reached highest concentration in the 0 to 3 cm layer, acetate at 3 to 6 cm and propionate at 6 to 9 cm. The results are discussed in relation to the role of these acids as food sources and as indicators of biochemical pathways taking place in sediments with different carbon input levels.     

Respiration and excretion by the ctenophore Mnepiopsis leidyi

Respiration (dissolved oxygen and carbon dioxide) and excretion (dissolved organic carbon, inorganic and organic nitrogen and phosphorus) rates were measured for a variety of sizes of Mnemiopsis leidyi over a temperature range of 10.3° to 24.5°C. Both respiration and excretion rates were a direct linear function of animal weight and very temperature sensitive (Q₁₀4). Oxygen uptake ranged from 155 to 489 g at O/(g dry weight) day^-1 and carbon dioxide release from 43 to 166 M. Organic carbon made up about 38% of the total carbon released. Inorganic nitrogen excretion, exclusively in the form of ammonium, comprised 54% of the total nitrogen release and ranged from 10 to 36 M NH₄/(g dry weight) day^-1. Average release of dissolved primary amines (expressed as glycine equivalents) equaled 43% of the organic nitrogen fraction. Inorganic phosphorus release ranged from 2.0 to 4.9 M/(g dry weight) day^-1 and made up about 72% of the total phosphorus loss. The turnover of elements in the body was calculated as 5 to 19% per day for carbon and nitrogen, depending on the temperature, and an even higher 20 to 48% per day for phosphorus. These values are comparable to rates observed for small, active zooplankton.     

Pattern of benthic biomass size spectra from shallow waters in the East China Seas

Benthic biomass size spectra (BSS) and normalized biomass size spectra were constructed, and benthic secondary production was estimated by a size spectrum equation in the shallow waters in the East China Sea, ranging latitudinally from 40°N to 29°N. The BSS patterns were bimodal, two biomass peaks corresponding to meiofauna and macrofauna, respectively, separated by a trough of low biomass at 8–256 μg individual dry weight which varied in position with median sediment particle size. The BSS also displayed bimodality within meiofauna size ranges, which in most stations was due to the relative proportions of nematodes and other meiofauna taxa. Re-analysis of data from sites in the UK, South Africa, and Antarctic showed a similar bimodality in the adult species body size distribution within the meiofauna size range. Macrofaunal production estimated by the size spectrum equation was very similar to the results of Brey90 empirical equation. However, these production values were much lower than those calculated by Brey01. Different individual dry-to-wet conversion ratios, temperature deviation, and macrofauna taxonomic composition might be responsible for the between-model differences. The macrofaunal P/B ratios calculated by this equation ranged from 0.3 to 3.4 which were in accordance with values from Northern Hemisphere mid-latitudes. Meiofaunal production estimates will need further empirical support.     

Ein neuer „Meiofaunastecher” zur quantitativen Probennahme in Weichböden

During 1972 and 1973, quantitative meiofauna samples were taken by subsampling out of a bottom sampler from some parts of the Kiel Bight. A new method has been developed to operate in very soft and muddy sediments with high organic and water contents. The construction of the new instrument is described. It consists of a bottomless plexiglass box, the cover and front of which may easily be removed. It does not cause compression of the sampled sediment, and hardly disturbs it. The semi-liquid surface of the sediment is kept in place by the use of a “sledge” and a strip of cellophane tape inside the sampler. Some preliminary results from one station show a meiofauna maximum (mainly nematodes) in the subsurface layer of the sediment, but this varied from one month to the next. The water content of the sediment at another station was as high as 300%.     

A simple density separation technique for quantitative isolation of meiobenthos using the colloidal silica Ludox-TM

A simple, quantitative density separation method is described. The method is based on differences in specific weight between meiobenthos and sediment. Nematodes and copepods could be separated from sediment and detritus when samples were suspended in Ludox-TM, a colloidal silica. Organisms float at the surface, while sediment particles sink. Results obtained with this new method were compared with the well-known decantation method. For a quantitative isolation of nematodes from sediments, rich in coarse detritus, a maximum volume of 7 cm³ sample could be used. For copepods this maximum was 13 cm³. For such sediments the density method is more reliable than the decantation method. The time needed for sorting the meiobenthic organisms is reduced to about 30% compared with the former method. The new method can be used for preserved as well as for fresh sediment samples and can also be applied for the disolation of small polychaetes, small oligochaetes, larvae of some macrofaunal groups and net-zooplankton.Publication No. 10 of the project Biological Research in the Ems-Dollard Estuary.     

Relationships between microbial distributions and the anaerobic decomposition of organic matter in surface sediments of long Island Sound,USA

Relative rates of the anaerobic decomposition of organic matter in the upper 10 cm of sediment from two stations in central Long Island Sound, USA, were compared. Sediment samples from discrete depth intervas were incubated anoxically and changes in SO ₄ ⁼ , NH ₄ ⁺ , bacterial numbers, extractable adenosine triphosphate (ATP), organic matter, and organic carbon were measured as a function of time and temperature. At both stations (15 and 34 m water-depths, respectively), the calculated rates of SO ₄ ⁼ reduction and NH ₄ ⁺ production decreased exponentially (approximately) with depth below the sediment-water interface. Over the same depth interval, ATP concentrations dropped by a factor of 6 to 7 and bacterial numbers were lower by a factor of 2 to 3. These decreases in SO ₄ ⁼ reduction, NH ₄ ⁺ production, bacterial numbers, and ATP, reflect a change in the physiological state of microbial populations with depth in the sediment and are consistent with the conclusions that the quantity of easily utilizable organic matter changes rapidly below the sediment surface and that food limitation controls the basic depth distribution of microbial activity. The average rates of SO ₄ ⁼ reduction, 29 to 39 mM year^-1 (22°C), in the top 10 cm are similar at both stations studied here, as well as at an additional station from a previous study. In contrast, average NH ₄ ⁺ production differs by a factor of 2 at the two stations, reflecting differences in the C:N ratio of the organic matter supplied to the sediment surface and differences in particle reworking by macrofauna at each site. The apparent activation energy of SO ₄ ⁼ reduction was 19±1 kcal mole^-1 and that of NH ₄ ⁺ production, 18±3 kcal mole^-1. The overall quantity of carbon required to support the calculated average SO ₄ ⁼ reduction rate in the top 10 cm is 23 g C m^-2 year^-1 and represents 36% of all the carbon available to the benthos annually and 11% of the net primary production in the water column. Directly measured fluxes of NH ₄ ⁺ from sediments to overlying water at both stations agree well with those predicted from production rates obtained by the incubation techniques.     

Daily ration and specific daily ration of the chaetognath Sagitta enflata

Digestion times, obtained directly from laboratory feeding, were used in conjunction with a day/night series of field samples to estimate the daily ration and specific daily ration of Sagitta enflata Grassi, the most abundant chaetognath in the Gulf Stream near Miami, Florida (USA), during winter and early spring. Feeding was independent of time of day and increased with increasing chaetognath size from 1.7 prey day^-1 for the 11.5 mm size class to 2.9 prey day^-1 for the 21.5 mm class. The diet of s. enflata consisted of 94.8% copepods having a mean weight of 21.8 g, and 5.2% prey chaetognaths estimated to be two-thirds the length of the predators. The daily ration of S. enflata from 12.5 to 20.5 mm in length increased from 48.3 to 143.3 g of prey dry weight and from 18.5 to 48.1 g of prey carbon. The specific daily ration declined from 0.12 to 0.08 dry weight basis, and from 0.26 to 0.14, carbon basis, over the same size range. Prey chaetognaths contributed as much as 51% of the ration on a carbon basis.