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)     

Observations of heterotrophic activity on photoassimilated organic matter

The natural heterotrophic utilization of released and particulate photoassimilated compounds by natural phytoplankton populations was followed during the summer of 1975 in the Kiel Bight (Western Baltic Sea). Phytoplankton exudates (2 to 21% of primary production) seem to represent an important substrate for heterotrophic bacteria: uptake rates between 8 and 17,5%/h were observed. Samples incubated with antibiotics indicated that the heterotrophic uptake of phytoplankton exudates during photosynthesis could lead to underestimation of the release rate. Production of 14_{CO 2}was used as an estimate of the mineralization of particulate photoassimilated matter. The mineralization rates measured were in the range of 6 to 20% after 20 h. In order to confirm the quantitative measurements, micro-autoradiographic examinations of the samples were performed.     

Nutrition of the temperate Australian soft coral Capnella gaboensis

We examined the ability of Capnella gaboensis Verseveldt, 1977 (Coelenterata: Octocorallia: Alcyonacea: Nephtheidae) to utilize heterotrophic food sources, and the importance of heterotrophic nutrition and photosynthesis in its diet, by using preserved material and histological sections of field-collected specimens and by means of laboratory experiments in which coral branches were fed with ¹⁴C-labelled food of different sizes. The study was conducted from April 1982 to August 1984. C. gaboensis receives nutrition from the photosynthesis of its symbiotic zooxanthellae, Symbiodinium sp., and from heterotrophic sources. Up to 10% of the algal photosynthate was translocated to the animal-host tissues. The contribution of translocated carbon from the zooxanthellae to the daily respiratory carbon requirement of the animal was estimated to be well below 50% in all seasons except in the summer of 1983–1984, indicating that the coral must rely on additional sources of nutrition (i.e., heterotrophy) for most, if not all, of the year. Field (Sydney Harbour: 33°50S; 151°15E) and laboratory observations and experiments indicated that this coral probably feeds upon zooplankton, small particulate matter and dissolved organic matter.     

Sedimentation of particulate matter during a phytoplankton spring bloom in relation to the hydrographical regime

Data presented and discussed here were collected continuously during April/May 1975 in the Bornholm Basin of the Baltic Sea. Sedimentation rates of particulate matter were recorded with 5 multisample sediment traps from different depths in the water column at 2 positions 170 km apart. Current meter data collected during the same period and depths indicated that the positions remained hydrographically distinct during the investigation. Particulate matter from the euphotic zone including diatom cells formed the bulk of the material collected by all traps. This flux of organic particles to the bottom was unimpeded by the strong density stratification present in the water column. The upper traps always collected less material than lower ones. This paradox has been ascribed to diminishing current speeds with depth, concomitant with an increase in sinking rates of phytoplankton and phytodetritus. Both factors influence the sampling efficiency of sediment traps, which are thought to have underestimated actual sedimentation rates here. A time lag of 2 to 3 weeks in bloom development seemed responsible for the characteristic differences between the two positions. The phase of major sedimentation at one position covered about 18 days, and a distinct sequence in the composition of the material collected by the 6 glasses of each trap indicated phases of a progressively deteriorating phytoplankton population in the water column contributing the particulate material. A total of 6.2 g C m^-2 in 34 days was recorded at this station. Apart from a trap situated in an oxygen deficient layer which collected 0.44 g C m^-2 of zooplankton corpses, zooplankton mortality was overestimated by the traps. Large-scale sedimencation of fresh organic matter produced by the spring bloom is probably a regular feature in areas with low over-wintering zooplankton populations and, as such, possibly has a direct stimulatory effect on growth and reproduction of the benthos.Contribution No. 185 of the Joint Research Programme 95, Kiel University.     

Some experiments on phosphate assimilation by coastal marine plankton

A study of phosphate assimilation by coastal marine plankton revealed that both phytoplankton and microheterotrophs incorporated radioactive phosphorus (³³P). Size fractionation of the particulate matter (using 1 m pore diameter Nucleopore^® membrane filters), antibiotic treatment (using garamycin), and independent estimaties of photoautotrophic (¹⁴CO₂ uptake) and heterotrophic (³H-glucose uptake) activities were employed to separate phyto- and bacterioplankton phosphate uptake. Results indicated that phytoplankton ³³P-uptake was best estimated by the fraction of particulate matter retained on the 1 m membrane filters. Usually, less than 10% of the phytoplankton (based on chlorophyll a measurements) passed the 1 m pore-diameter filters, whereas about 90% of the heterotrophic activity passed. At least 50% of the ³³P-uptake was associated with the <1 m fraction. It may be possible to resolve the phytoplankton and bacterial contributions to ³³P-uptake by comparing the percent of total ³³P-uptake with the percent of total ³H-glucose uptake associated with the >1 m fraction.     

Relationships between marine microorganisms and the wood-boring isopod Limnoria tripunctata

The quantities and the behavior of particulate proteinous carbon (PPC) were investigated at an experimental station in Funka Bay, southern Hokkaido from February 1974 to January 1975 using R. V. Ushio Maru. The amino acid composition of PPC, quantity of particulate organic carbon (POC), PPC:POC ratio, concentration of plant pigments and the primary productivity and heterotrophic activity in situ were measured to clarify PPC status. The net production was ca. 100 g C m^-2 year^-1. The inflow of Oyashio water and stagnation during early spring and summer may increase productivity. The net production during this period attained 70% of the net production for the whole year. The concentration of chlorophyll changed monthly parallel to changes in the primary production of the surface waters. The concentration of PPC and the PPC:POC ratio in the surface layer decreased gradually during the stagnation period of the Oyashio water. This implies that the PPC in the particulate organic matter was more easily consumed than the other organic constituents by the increased heterotrophic activity in summer. In contrast, low concentrations of PPC and high PPC:POC ratios were found during the stagnation period of Tsugaru warm water from autumn to winter. Concentrations of PPC and POC were not evenly distributed throughout the year; they decreased with depth during the stagnation period in summer. On the other hand, higher concentrations of PPC and POC, and an increased PPC:POC ratio were noted in samples collected near the sea bed, compared with the concentration and the ratio of the upper layer throughout the year. It seems likely that the particulate organic matter, originating from resuspension of bottom sediments containing proteins, was produced by the benthos. The turnover time of PPC in the upper layer was 7 to 11 days in summer and 45 to 100 days in winter; the turnover time for POC was 14 to 28 days in summer and 97 to 147 days in winter.     

Zooplankton abundance and grazing at Davies Reef,Great Barrier Reef,Australia

Zooplankton abundance and grazing on autotrophic and heterotrophic particulate matter were measured along a transect across Davis Reef (18°5S; 147°39E) and in the back-reef lagoon over tidal and diel cycles during austral winter (August 1984). Zooplankton entering the reef from the surrounding shelf waters decreased in abundance over the reef flat, presumably because of predation. Within the reef lagoon, maximum daytime densities of pelagic copepods occurred during high water, suggesting an external input. At night, water-column zooplankton biomass increased by a factor of 2 to 3 due to the emergence of demersal reef zooplankton. Zooplankton grazing rates on heterotrophic particulate matter (bacteria + detritus and Protozoa) compared to phytoplankton were higher on the reef flat than on the fore-reef or lagoon. Within the lagoon, zooplankton grazing rates on heterotrophic material were maximum during high water, coincident with maximum tidal concentrations of particulate organic carbon. The combined demersal and pelagic zooplankton community were often able to crop 30% of the daily primary production by >2µm phytoplankton. However, >50% of phytoplankton biomass was in cells <2µm, presumably unavailable to these zooplankton. Our particulate production and ingestion measurements, together with zooplankton carbon demand extrapolated from respiration estimates, suggest that the zooplankton community of Davies Reef derives much of its nutrition from detritus.Joint contribution from the University of Maryland, Center for Environmental and Estuarine Studies (No. 2015), and the Microbial Ecology on a Coral Reef Workshop (MECOR No. 19)     

Laboratory study of effects of temperature and salinity on survival and larval development of a population of Rhithropanopeus harrisii from the Mondego River estuary,Portugal

Rhithropanopeus harrisii (Gould) is an introduced species in the estuary of the Mondego River (Portugal): it was first recorded from the Iberian Peninsula in 1989. The larval development of this population was studied under laboratory conditions at different temperatures and salinities, and showed a larval development pattern very similar to that reported for American estuarine populations of this species. Larval development was negatively correlated with temperature. Time to megalopa varied between 7 and 35 d; the first crab (C₁) was reached after a maximum of 11 to 43 d. Larval development was optimum at 25°C and 15S. Larval survival was maximum at 10, 15 and 20S at all three temperatures studied (20, 25 and 30°C). The percentage of abnormal megalopae increased with increasing salinity to a maximum (100%) at 30S; incidence of abnormality was not affected by temperature.     

Comparison of the sedimentation of a diatom spring bloom and of a subsurface chlorophyll maximum

The seasonal pattern of sedimentation was determined over a 8-mo investigation period covering the productive season at a permanent station in the Southern Kattegat (Denmark) in 1990. The phytoplankton succession was characterised by a 4-mo long subsurface maximum associated with the pycnocline which was entirely dominated by the dinoflagellate Gyrodinium aureolum. The bulk sedimentation of organic matter took place during this period and accounted for >60% of the annual particulate organic carbon (POC) and particulate organic nitrogen (PON) sedimentation. The spring bloom period contributed 60% of the sedimentation of intact phytoplankton cells, but only 20% of the POC and PON sedimentation. A minor fraction of the sedimenting matter from the subsurface phytoplankton maximum consisted of intact phytoplankton (<20%), suggesting that the phytoplankton was processed by heterotrophs and that it was mainly products from this activity which contributed to thevertical flux of organic matter. The variation in oxygen concentration below the pycnocline coincided with the pattern of sedimentation with a delay of 3 to 6 wk.     

The heterotrophic phase of plankton succession in the Japan Sea

The vertical structure, composition and productivity of a plankton community was studied in the Japan Sea in June, 1972 during a period of thermocline formation; the parameters measured were: phytoplankton production and biomass; number, biomass, and production of planktonic bacteria; biomass of phagotrophic flagellates, ciliates and remaining microzooplankton. The concentration of micro- and mesozooplankton attained a basic maximum in a layer near the upper part of the thermocline. The biomass and calculated production of the heterotrophic part of the community exceeded considerably the amount of primary production. The heterotrophic phase of the seasonal succession of a plankton community in a temperate sea is described, when heterotrophic metabolism and production predominate. Heterotrophs at this stage use mostly energy from organic matter accumulated during the previous spring phytoplankton bloom.     

Effect of temperature and salinity on larval development of southern king crab (Lithodes antarcticus)

Larvae of Lithodes antarcticus Jacquinot were reared in October, 1981 from hatching to the glaucothoe stage at 16 temperature/salinity combinations (5.5°; 7.5°; 9.5° and 13.5°C; 26, 29, 32 and 35 S) to determine optimal environmental conditions for larval development. The highest survival percentage was obtained in the culture at 7.5°C and diminished according to temperature increase or decrease. High temperature cultures significantly shorten the larval life duration, but produce large mortalities. At 5.5°C mortality occurred almost exclusively during the moult to glaucothoe stage. Higher survival percentages were obtained as salinity was increased. In the lowest salinity culture (26 S) no zoea reached the post-larvae stage at culture temperatures. The best T/S combination was obtained at 7.5°C and 35 S, with a survival percentage of 29%. The shortest zoeal developments were obtained at 32 S in all culture temperatures. Salinity also affects larvae coloration: there is a pigment concentration on erythrophores, which causes a color decrease.     

Distribution of labile and residual particulate carbon in the Baltic Sea

During the September 1971 cruise of F. K. Alkor in the central Baltic Sea, the surface or summer water layer down to a depth of about 30 m was found to contain 187.4 g/l of particulate organic carbon, with a C:N (atoms) ratio of 8.97. This carbon was 44% (89.9 g/l) labile to bacterial decomposition, as determined by burning an aliquot of each sample in a CHN-analyser before and after maintenance in nutrientenriched sea water at 20°C for 3 months. The particulate material from the intermediate or winter water layer, with a depth ranging from 30 to 70 m, contained only 48% (43.1 g/l) of the labile carbon found in the summer surface layer, and had a significantly higher C:N ratio (11.25). These two facts indicate that a considerable breakdown of the organic material had taken place. The material, removed from the particulate state during this process, was 48% (89.7 g/l) of the original total particulate carbon, and was relatively nitrogen-rich, with a C:N ratio of 6.49. In this material, 52% (46.8 g/l) of the organic carbon was labile. The particulate material in the rest of the water column showed no significant changes until it reached the sediment. The slurry immediately above the sediment had a C:N ratio of 9.15, indicating the introduction of nitrogen from either dissolved or colloidal material. Labile carbon (44% of the total) was also present in sufficient quantities to support life and to make this an important diagenetic site.     

Calcification in the maerl coralline alga Phymatolithon calcareum: Effects of salinity and temperature

The coralline alga Phymatolithon calcareum was dredged from 13 m in the Kattegatt, Baltic Sea, in December, 1980, and its rate of calcification was measured by ⁴⁵Ca⁺⁺-uptake methods. Light-saturated calcification rates at 5°C ranged from 15.8 g CaCO₃ g^-1 dry wt h^-1 for the basal parts of the plants to 38.7 g CaCO₃ g^-1 dry wt h^-1 for the tips. These age gradients were not apparent when calcification rates were expressed on the basis of surface area. Experiments with salinity (10, 20, 30) and temperature (0°, 5°, 10°, 20°C) indicated that optimum conditions for calcification were at 30 S and at temperatures above 10°C. Salinity had a greater influence on calcification rate than did temperature, and there was a positive relationship between salinity and calcification rate at all temperatures. In 6 mo old cultures, salinity was again the important factor, with all plants remaining healthy at 30 except those at the highest temperature (20°C). These trends, and the low calcification rates at 10S (4.6 g CaCO₃ g^-1 dry wt h^-1 at 5°C to 8.6 g CaCO₃g^-1 dry wt h^-1 at 20°C) suggest that low salinity may be the explanation for the general absence of P. calcareum from the brackish waters of the Baltic Sea. Short-term experiments in which salinity was kept constant while Ca⁺⁺ concentration was altered, and experiments in which salinity was varied and Ca⁺⁺ concentration kept constant, suggest that it is the calcium ion concentration and not salinity per se which affects calcification rates.     

Production,composition and sedimentation of salp fecal pellets in oceanic waters

Rates of fecal pellet production have been recorded from seven species of oceanic salps feeding on natural diets. Expressed as g C defecated per mg salp body C per hour, the values range between 3.7 and 27.7. Carbon: nitrogen ratios of the salp fecal pellets average 11.4; the organic matter of the pellets is mainly protein and carbohydrate. Sinking velocities of the pellets are very high, ranging from 320 to 2 238 m d^-1 for pellets from three species. However, the pellets sink slower than would be predicted from extrapolation of rates for crustacean pellets, probably due to the shape of the pellets and their density. The high rates of defecation, large size and rapid sedimentation of salp fecal pellets make them likely mechanisms for rapid transport of small particulate matter from surface waters to deep water and the benthos.     

Molecular divergence between North Atlantic and Indo-West Pacific Cladophora albida (Cladophorales: Chlorophyta) isolates as indicated by DNA-DNA hybridization

Genomic relationships between North Atlantic, Australian and Japanese isolates of the benthic seaweed Cladophora albida (Huds.) Kütz. were examined in 1987 by means of DNA-DNA hybridization. The data indicate that C. albida can be divided into a North Atlantic and an Indo-West Pacific group with an intergroup hybridization response of 25 to 30% and 5.5° to 6.0°C for hybridization percentage and T _me the melting temperature reduction of hybridized sequences, respectively. This level of genome divergence is considerably higher than that observed in most other eukaryotes. The separation between the two C. albida groups presumably dates back to the closure of the Asian part of the Tethys Ocean, about 12 million years ago. The data also indicate that transatlantic C. albida populations have a greater genetic inter-relatedness than have Japanese and Australian populations. In C. albida there is no clear correlation between molecular evolution and the evolution of morphological traits. C. albida and C. rupestris (L.) Kütz have hardly any DNA sequences in common.     

Effects of temperature and salinity on larval development ofNecora puber (Brachyura: Portunidae)

Temperature and salinity affected both length of larval development and mortality inNecora puber collected in the Ría de A Coruña during December 1984 and January 1985. Development time decreased considerably with increased temperature. This decrease was sharper when temperature increased from 15° to 20°C than when it increased from 20° to 25°C. At 35S, average development took 48, 32 and 28 d at 15°, 20° and 25°C, respectively. At the three salinities tested (25, 30 and 35), larval development was completed only at 15°C, at 20°C/30 and 35S, and at 25°C/35S. Development times at 15° and 20°C were highly significantly different at both 35 and 30S (P 0.01). However, there were no significant differences between development times at 20° and 25°C (P > 0.05). Within any one specific temperature series, no significant difference was observed between the salinity values tested (P > 0.05). The duration of each of the five zoeal stages was similar within each and the same temperature/salinity combination, whereas the duration of the megalop was twice as long as any of the zoeal stages. The combination of the lowest temperature (15°C) and the highest salinity (35) tested resulted in the greatest larval survival of 28%. Highest mortality occurred at 25°C, at which temperature development was completed only at 35S. A sharp drop in larval survival was observed in the transition period Zoea V — megalop in all combinations of temperature and salinity tested. Within the limits of tolerance to temperature and salinity, the former effected more pronounced differences in the duration of larval development, while salinity appeared to constitute a limiting factor for survival.     

Metabolic diversity in epibiotic microflora associated with the Pompeii wormsAlvinella pompejana andA. caudata (Polychaetae: Annelida) from deep-sea hydrothermal vents

Specimens of alvinellid polychaetes (Alvinella pompejana Desbruyères and Laubier, 1980 andA. caudata Desbruyères and Laubier, 1986) and their tubes were sampled from deep-sea hydrothermal vents at 13°N from the manned submersible Nautile during the Hydronaut cruise (October to November 1987) on the East Pacific Rise. Samples were subjected to bacterial analysis aboard the mother ship Nadir to detect bacteria involved in the nitrogen and sulphur cycles, in non-specific heterotrophic processes, and displaying resistance to selected heavy metals. Cultures were incubated at different temperatures under atmospheric and in situ (250 atm) pressures. Bacterial growth was observed in enrichment cultures for most metabolic types screened. Heavy-metal-resistant bacteria were also detected in many samples. No filamentous bacterial form was observed in the cultures. The results demonstrate a high metabolic diversity in episymbiotic flora, indicating that the worm (A. pompejana orA. caudata), its tube and its epiflora represent a complex micro-ecosystem.     

Croissance et structure de l'opercule calcifié du gastéropode polynésien Turbo setosus (Prosobranchia: Turbinidae): détermination de l'âge individuel

Turbo setosus Gmelin were collected in 1979 and 1982 in French Polynesia (Tuamotu). Ground cross-sections of calcified opercula revealed 3 distinct types of growth lines (A, B and C), the appearance and periodicity of which are described. Two months after fluorescent labelling with alizarin and fluorescein, permanent marks were easily visible in the operculum, enabling the 3 types of growth lines to be interpreted and a growth pattern distinguished: Type A growth lines, which are very translucent under transmitted light, are irregularly deposited after individual traumatisms and are termed stress lines; Type B lines exhibit a regular periodicity and correspond to a daily growth rhythm; Type C lines reflect a subdaily growth pattern. Vital staining enabled calcification of the operculum to be followed, and the speed of deposition has been estimated. The textural organization of the organic and mineral phases varies according to operculum area and depends on local speed of growth. In those areas of rapid growth (more than 20 m d^-1), deposits are cone-shaped and interlocked, and in those areas where growth is slower (less than 20 m d^-1), this type of organization disappears and is replaced by a regular organization without cone-shaped deposits. The number of daily growth lines, counted on 66 opercula, allowed us to accurately assess individual age and longevity in this species, which did not exceed 3 yr in our sample.     

Size composition of particulate organic matter in the lagoon of Tikehau atoll (Tuamotu archipelago)

Suspended particulate matter was comprehensively investigated from 6 to 17 April 1986 in the lagoon of Tikehau atoll (15°00S; 148°10W). Dry weight (DW), particulate organic carbon (POC), adenosine triphosphate (ATP), and chlorophyll a were measured for five size-classes (0.2 to 0.8 m, 0.8 to 3 m, 3 to 35 m, 35 to 200 m, and 200 to 2000 m). Taxa were identified and counted for the whole plankton (both autotrophic and heterotrophic). Particles <3 m accounted for 81% of the total POC (192 mg m^-3), and detritus comprised 82% of the total POM. Phytoplankton (cyanobacteria plus algae) accounted for 35% of the living carbon, 75% of which consisted of heterotrophic bacteria and cyanobacteria. The zooplankton biomass was composed of 31% nano-, 26% micro-, and 43% mesoplankton.     

Effect of growth factors and light quality on the growth,pigmentation and photosynthesis of two diatoms,Thalassiosira gravida and Phaeodactylum tricornutum

Recruitment, life span and growth rate were investigated in field and experimental populations of Abra alba (Wood) in Kiel Bay, FRG (55°N) from 1975 to 1978 to determine production to biomass (P:B) ratios and to assess the importance of A. alba to production by commercial fish. Life span and growth rates were determined from changes in length frequency modes at each site and from winter rings on the shell. A peak of recruitment usually occurred in August, sometimes followed by a second peak between December and February. Life span was between a little more than one year and two and a half years. Growth rates were highest at the two sites in offshore fishing grounds, where bivalves reached a mean length of 13 to 16 mm at the end of two years. At the inshore control site and in the nearby experimental containers, individuals reached a mean length of 7 mm at the end of two years. Production estimates ranged from 110 to 3.000 mg C m^-2 year^-1, differing markedly among sites and among years. Production was highest during the first year after recruitment, occurring mainly between July and December. Mortality occurred mainly between January and June, and was in approximate balance with production over a three-year period. Annual P:B ratios were from 1.3 to 3.4; a long-term mean P:B ratio of 2.2 is suggested for Kiel Bay populations of A. alba. Annual production by A. alba appears to exceed considerably consumption by fish of commercial size. The significance of A. alba in the food web of Kiel Bay may thus be as food for juvenile fish or for intermediate-level predators that are themselves prey for larger fish.Publication No. 431 of the Sonderforschungsbereich 95 Meer-MeeresbodenPublication No. 1031 of the CSIRO Marine Laboratories