Excretion of ammonia by zooplankton and its potential contribution to nitrogen requirements for primary production in the Catalan Sea (NW Mediterranean)

Excretion of ammonia by mesozooplankton (>200 m zooplankton) and its potential contribution to the nitrogen requirement for phytoplankton growth has been estimated for different hydrographical situations along a transect across the Catalan Sea (Northwestern Mediterranean). The nitrogen excreted as ammonia was estimated from mesozooplankton biomass and specific excretion rates. Nitrogen requirements of phytoplankton were estimated by means of carbon fixation rates and C:N ratios of <200 m particulate organic matter. Minimum C:N ratios and maximum primary production, zooplankton biomass, phytoplankton nitrogen requirements, and nitrogen excretion of zooplankton occurred near the Catalan density front. On average, the nitrogen regenerated by the mesozooplankton accounted for 43% of the nitrogen requirements of the phytoplankton. The specific excretion rates of ammonia and the percentage of phytoplanktonnitrogen requirements supplied by excreted nitrogen were higher at coastal stations. In some coastal and frontal stations, the ammonia excreted exceeded the phytoplanktonnitrogen demand. Bacteria competing for nutrient supply and the possible uncoupling between rate processes and standing stocks of phyto- and zooplankton could explain the apparent excess of regenerated ammonia.     

Succession and growth limitation of phytoplankton in the Gulf of Bothnia (Baltic Sea)

A one year field study of four stations in the Gulf of Bothnia during 1991 showed that the biomass was ca. two times, and primary productivity ca. four times, lower in the north (Bothnian Bay) than in the south (Bothnian Sea) during the summer. Nutrient addition experiments indicated phosphorus limitation of phytoplankton in the Bothanian Bay and the coastal areas in the northern Bothnian Sea, but nitrogen limitation in the open Bothanian Sea. A positive correlation between the phosphate concentration and the production/biomass ratio of phytoplankton was demonstrated, which partly explained the differences in the specific growth rate of the phytoplankton during the summer. Differences in photosynthetic active radiation between the stations also showed a covariation with the primary productivity. The relative importance of nutrient or light limitation for photosynthetic carbon fixation could not, however, the conclusively determined from this study. Marked differences in phytoplankton species composition from north to south were also observed. The number of dominating species was higher in the Bothnian Sea than in the Bothnian Bay. The distribution of some species could be explained as due to nutrient availability (e.g. Nodularia spumigena, Aphanizomenon sp.), while salinity probably limits the distribution of some limnic as well as marine species. The potentially toxic phytoplankton N. spumigena, Dinophysis acuminata and Chrysochromulina spp. were common in the Bothnian Sea but not in the Bothnian Bay. The pico- and nanoplankton biomass during late summer was higher than previously reported due to a revised carbon/volume ratio.     

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.     

Production and respiration of overall plankton and ultraplankton communities at the entrance to the Gulf of Finland in the Baltic Sea

Primary productivity and respiration of the overall plankton community and of ultraplankton (organisms passing through a 3-m Nuclepore filter) were studied at the entrance to the Gulf of Finland during the growth season in 1982. Data of the respiration measurements from previous years are also presented. During the development of a diatom spring bloom, the algal component could be successfully separated from the bacterial component by size fractionation with a 3-m Nuclepore filter and thus the algal respiration could be approximated, being on the order of 10 to 20% of the gross production. After the phytoplankton spring maximum, bacteria played an important role in mediating the energy flow from phytoplankton exudates to higher trophic levels. Maximum values of 1 230 and 740 mg O₂ m^-2 d^-1 were recorded for overall and for ultraplankton respiration, respectively, during late July. High productivity values coupled with low phytoplankton biomass and low inorganic nutrient values were also recorded in late July, indicating effective nutrient regeneration and rapid turnover of the plankton community. During late summer, a considerable fraction (over 30%) of phytoplankton production was released as exudates, suggesting that much of the energy is channeled to higher trophic levels via bacterial pathways rather than by direct herbivorous grazing during this season. The summer development of phytoplankton community structure and functioning is strongly controlled by hydrographic conditions, i.e. by nutrient inputs via upwelling and by water temperature. A carbon budget for late summer indicated that bacteria may contribute only up to 50% of the overall respiration of the plankton community, which suggests that heterotrophs other than bacteria play an important role in nutrient regeneration. The present study stresses the importance of energy flow via the phytoplankton exudatebacteria-micrograzer pathway in relatively oligotrophic, brackish water ecosystems.     

Net-zooplankton biomass of the Adriatic Sea

On the basis of 1 107 plankton samples collected during 1971–1981 in the different parts of the Adriatic Sea, the horizontal distribution of net zooplankton biomass, expressed as dry weight and ash-free dry weight, is discussed. The northern Adriatic combined with the Gulf of Trieste usually had the highest standing crop of zooplankton. Biomass in other regions was lower and more uniform. In general, the highest zooplankton stocks came from the shallow regions close to the coast where the sources of enrichment from the land (fresh water inflow, land drainage. pollution where it exists) and the shallow sediments are the highest. We believe the elevated zooplankton biomass encountered in the north Adriatic to be due to proliferation of tolerant non-selective feeding zooplankton species, such as copepods, which can use detritus as a major fraction of their diet. The northern Adriatic is rich in living and non-living suspended particles throughout the year and can thus maintain high zooplankton biomass. This conclusion is probably valid for similar regions of the Mediterranean Sea.     

Microbial hydrolysis of polysaccharides and organic phosphates in the Northern Adriatic Sea

g -glucosidase and alkaline phosphatase, two ectoenzymes involved in the microbial decomposition of polysaccharides and organic phosphates, were evaluated in water samples collected from two areas of the Northern Adriatic Sea during the multidisciplinary Prisma II research project. The distribution of g -glucosidase and alkaline phosphatase is reported together with that of leucine aminopeptidase, an enzyme involved in the degradation of protein compounds. The data obtained showed the prevalence of g -glucosidase in summer months in the northern area, located in front of the Po delta, while in winter it is higher in the southern area (in front of Ancona). Phosphatase activity during February, 1998 had a maximum of 107 r nml m 1 h m 1 at a coastal station in the northern area; this enzyme appeared to be mainly associated with phytoplankton because the increase in alkaline phosphatase was associated with the increase of phytoplankton biomass, measured by chlorophyll a content. The relations between the microbial activities and environmental parameters are discussed.     

Phytoplankton photosynthetic activity and growth rates in the NW Adriatic Sea

Taxonomic composition, biomass, primary production and growth rates of the phytoplankton community were studied in two stations in the NW Adriatic Sea on a seasonal basis, in areas characterized by differing hydrological and trophic conditions. The main differences between the two stations were quantitative rather than qualitative, most phytoplankton species being common to both stations. The effects of differing nutrient concentrations and plume spreading were evident. Biomass and primary production rates were significantly higher in the coastal station (S1), and the phytoplankton distribution in the water column was markedly stratified in S1 and more even in the offshore station (S3). However, chlorophyll a specific production, potential growth rate and production efficiencies were very similar in both stations, even when phosphorus concentrations were limiting. A discrepancy between potential and actual growth rate was observed: as a feature common to both stations, comparisons between potential and actual growth rates revealed that little carbon produced by phytoplankton accumulated as algal biomass; therefore, very high loss rates were estimated.     

Phytoplankton photosynthetic activity and growth rates in the NW Adriatic Sea

Taxonomic composition, biomass, primary production and growth rates of the phytoplankton community were studied in two stations in the NW Adriatic Sea on a seasonal basis, in areas characterized by differing hydrological and trophic conditions. The main differences between the two stations were quantitative rather than qualitative, most phytoplankton species being common to both stations. The effects of differing nutrient concentrations and plume spreading were evident. Biomass and primary production rates were significantly higher in the coastal station (S1), and the phytoplankton distribution in the water column was markedly stratified in S1 and more even in the offshore station (S3). However, chlorophyll a specific production, potential growth rate and production efficiencies were very similar in both stations, even when phosphorus concentrations were limiting. A discrepancy between potential and actual growth rate was observed: as a feature common to both stations, comparisons between potential and actual growth rates revealed that little carbon produced by phytoplankton accumulated as algal biomass; therefore, very high loss rates were estimated.     

Assessment of distubance in Mediterranean lagoons: An evaluation of methods

A number of methods were applied to assess disturbance in macrozoobenthic communities in three Mediterranean lagoons with different levels of stress. Tsopeli, Ionian Sea, with no obvious source of stress, harbours a fauna typical of brackish-water lagoons. Vivari, Aegean Sea, also without apparent source of disturbance, is characterised by a few typical lagoonal species and some species characteristic of perturbation. Goro lagoon, in the northern Adriatic, is much larger and more open to the sea. At the centre of the lagoon, where anoxia is known to occur in the summer (Goro polluted), the fauna is dominated by species typical of disturbance and a few lagoonal species. A dredged area closer to the sea (Goro dredged) is totally dominated by species characteristic of disturbance. The species diversity in all lagoons ranges from low to very low. According to the distribution of individuals in geometric abundance classes, all the lagoons are characterised as stressed. The distribution of individuals in geometric size classes shows dominance of larger specimens in the least disturbed Tsopeli and exclusively small sizes in the greatly disturbed dredged area of Goro. The abundance/biomass comparison curves characterise Tsopeli as undisturbed, Vivari and Goro polluted stations as moderately disturbed and Goro dredged station as disturbed. The last two methods agree with the characterisation derived by examining the dominant species. It is concluded that methods based on size changes of the fauna are more sensitive than those based on relative abundance in assessing disturbance in coastal brackish-water lagoons.     

Mechanisms of production and fate of organic phosphorus in the northern Adriatic Sea

In the period from 1980 to 1984 organic phosphorus, nutrients, primary production rates (¹⁴C), chlorophyll a (chl a) standing crops, and basic oceanographic parameters were measured during 23 cruises at six stations in the open waters of the northern Adriatic Sea. These waters are significantly influenced by polluted Po River discharge. Organic phosphorus was correlated with several parameters which characterize phytoplankton activity and organic matter decomposition processes. In the late winter-spring period, organic phosphorus is produced during phytoplankton blooms. It is hypothesized that microzooplankton grazing is the main factor increasing the organic phosphorus concentrations in summer (up to 1.1 mol 1^-1). Fall and winter had much lower values (below 0.3 mol 1^-1) due to remineralization processes and an increased water mass exchange between the northern and central Adriatic regions. The direct contribution of organic phosphorus by freshwater discharge was not found to be significant. The higher organic phosphorus concentrations that can occur in low salinity waters are most likely due to their increased capability to support primary production.     

Interactions between nutritional status and long-term responses to ultraviolet-B radiation stress in a marine diatom

Influences of nutritional status on the photoinhibitory effects of ultraviolet-B radiation (UVBR: 290 to 320 nm) on the specific growth rates (_obs) and biomass of Phaeodactylum tricornutum were determined using nutrient-replete batch cultures and nutrient-limited continuous cultures. P. tricornutum cultures were exposed to UVBR doses representative of current mid-latitude and ozone-depletion intensities. Specific growth rates and biomass were inhibited from 2 to 16% by UVBR during nutrient-replete growth. However, no effect of UVBR on _obs or biomass was detectable when nutrient limitation exceeded the potential for limitation by UVBR. Thus, a competitive interaction appears to occur between macronutrient stress and UVBR stress, such that _obs and biomass will be determined by the most limiting factor. Our results suggest that measurable decreases in phytoplankton _obs and biomass from UVBR are most likely in nutrient-rich areas of the ocean, while these parameters may not be appropriate for measuring UVBR stress in regions of nutrient limitation.     

Mesozooplankton community structure in offshore and coastal waters of the Ionian Sea (eastern Mediterranean) during mixed and stratified conditions

Depth-stratified samples, collected during a period where the water column was vertically mixed (March 2000) and a period of thermal stratification (September 2000), were analyzed in order to investigate the horizontal and vertical distribution patterns and composition of mesozooplankton, especially copepod species assemblages, in a pelagic (Ionian Sea) and a coastal area (Patraikos Gulf) of the eastern Mediterranean. Total mesozooplankton abundance and biomass were significantly lower in the highly oligotrophic offshore waters of the Ionian Sea when compared to the semi-enclosed Patraikos Gulf during both seasons. Small-sized copepods dominated the mesozooplankton community. An ‘offshore’ and a ‘coastal’ copepod assemblage were defined in the surface layer (0–50 m) only during March when differences in environmental conditions (i.e., temperature, salinity and fluorescence) were strong between the two areas. Copepod vertical community structure in offshore waters differed between sampling months. In March one assemblage (0–200 m) was mainly identified, while in September three distinct assemblages (0–50, 50–100 and 100–200 m) were observed, related to different vertical distribution patterns of the various copepod species. A pronounced seasonal change of the dominant copepods was evident in the surface layer, where strong differences in hydrological properties were observed from March to September. Below this layer, the copepod community was relatively stable showing decreasing seasonal differences with increasing depth.     

Climate change effects on phytoplankton depend on cell size and food web structure

We investigated the effects of warming on a natural phytoplankton community from the Baltic Sea, based on six mesocosm experiments conducted 2005–2009. We focused on differences in the dynamics of three phytoplankton size groups which are grazed to a variable extent by different zooplankton groups. While small-sized algae were mostly grazer-controlled, light and nutrient availability largely determined the growth of medium- and large-sized algae. Thus, the latter groups dominated at increased light levels. Warming increased mesozooplankton grazing on medium-sized algae, reducing their biomass. The biomass of small-sized algae was not affected by temperature, probably due to an interplay between indirect effects spreading through the food web. Thus, under the higher temperature and lower light levels anticipated for the next decades in the southern Baltic Sea, a higher share of smaller phytoplankton is expected. We conclude that considering the size structure of the phytoplankton community strongly improves the reliability of projections of climate change effects.     

Fine-scale distribution of juvenile Atlantic and Mediterranean loggerhead turtles (Caretta caretta) in the Mediterranean Sea

Loggerhead turtles nesting in the Mediterranean Sea exhibit remarkable genetic structuring. This paper tests the hypothesis that young loggerhead turtles from different rookeries do not distribute homogeneously among the major Mediterranean foraging grounds, due to a complex pattern of surface currents. We extracted long fragments of mitochondrial DNA from 275 stranded or bycaught juvenile turtles from six foraging grounds (Catalano-Balearic Sea, Algerian basin, Tyrrhenian Sea, Adriatic Sea, northern Ionian Sea and southern Levantine Sea). We used a Bayesian mixed-stock analysis to estimate the contributions from rookeries in the Mediterranean, the North-west Atlantic and Cape Verde to the studied foraging grounds. Differences were found in the relative contribution of juvenile turtles of Atlantic and Mediterranean origin to each foraging ground. A decreasing proportion of Atlantic juveniles was detected along the main surface current entering the Mediterranean, with a high prevalence of turtles from eastern Florida in the Algerian basin and lower numbers elsewhere. In regard to the turtles of Mediterranean origin, juveniles from Libya prevailed in central and western Mediterranean foraging grounds other than the Algerian basin. Conversely, the Adriatic Sea was characterised by a large presence of individuals from western Greece, while the southern Levantine Sea was inhabited by a heterogeneous mix of turtles from the eastern Mediterranean rookeries (Turkey, Lebanon and Israel). Overall, the distribution of juveniles may be related to surface circulation patterns in the Mediterranean and suggests that fisheries might have differential effects on each population depending on the overlap degree between foraging and fishing grounds.     

Phosphorus dynamics associated with phytoplankton blooms in eutrophic Mikawa Bay,Japan

Changes in the phosphorus components of the particulate matter in seawater were studied in the eutrophicated waters of Mikawa Bay, Japan, during summer 1981. The contents of particulate phosphorus and hot-water extractable intracellular phosphorus displayed remarkable changes associated with phytoplankton blooms caused by wind-induced or upwelling-associated nutrient enrichment from the bottom water layers. Nanoplankton <10 m accounted for much of the particulate phosphorus (70 to 79% in June and July, and 44 to 78% in August and September); the contribution of large-sized phytoplankton >25 m varied from 9 to 49%, the peak values being attained under red-tide conditions. The capacity for phosphorus storage in cells was low in nanoplankton cells, high in large phytoplankton species. Differences in rates of phosphorus storage and growth between nanoplankton and large phytoplankton accounted for fluctuations in particulate phosphorus which were closely associated with fluctuations in phytoplankton blooms in Mikaw Bay.     

Nutrient balance in the ecosystem of the North Western Adriatic Sea

The effects of biological processes on dissolved inorganic nutrients, dissolved organic nitrogen (DON) and phosphorus (DOP) are considered in the north western Adriatic Sea. The budgets of these nutrients, which represent the sum of production and consumption processes in comparison to advection, are discussed with regard to dissolved inorganic nitrogen ( 15 N labelled) uptake, which basically indicates the biological demand of this fraction of nitrogen by phytoplankton community. The presented data show that, although important, the continental input of dissolved inorganic nitrogen (DIN), mainly nitrate, is utilised and recycled within the coastal marine environment (budget of m 15 r µmol-N·dm m 3 ). In fact, during four cruises (June, 1996; February, 1997; June, 1997; February, 1998), phytoplankton production was mainly driven by regenerated nutrients ( f h 0.4). Regarding dissolved inorganic phosphorus (DIP), the negative budgets observed in most cases (down to m 0.4 r µmol-P·dm m 3 ) confirm, above all, its scarce availability in this basin. Recycling processes rather than continental inputs regulate the availability of this nutrient. In addition, the comparison between DIN and DIP budgets indicates that, in this ecosystem, dissolved inorganic phosphorus is recycled faster than nitrogen through the living particulate and dissolved organic pools. As a consequence of biological activities, a strong production of dissolved organic nitrogen (DON) can occur in summer (up to +22 r µmol-N·dm m 3 ) while DOP shows a more independent behaviour both with respect to its accumulation in the environment and to the observed nitrogen variations.     

The annual cycle of protozooplankton in the Kiel Bight

Protozooplankton (heterotrophic dinoflagellates and ciliates) composition and biomass was studied in a 20-m water column in the Kiel Bight on 44 occasions between January 1973 and April 1974. Both groups attained comparable biomass maxima during spring and autumn (0.3 to 0.7 g C m^-2 in the 20-m water column) and biomass levels were much lower in summer and lowest in winter. The spring protozooplankton maximum coincided with that of phytoplankton and during the rest of the year, protozooplankton stocks did not appear to be food limited as phytoplankton stocks were large throughout; many protozoans with ingested microplankton cells were observed, indicating that their potential food supply is not restricted to nanoplankton. Non-loricate organisms dominated biomass of the ciliates and tintinnids were of little importance. Tintinnids predominated in plankton samples concentrated by 20 m gauze indicating that most non-loricate ciliates, irrespective of size, were not retained. When phytoplankton sotcks were large (>3 g C m^-2) but those of metazooplankton small, as in spring and autumn, protozooplankton were the major herbivores with biomass levels comparable to those attained in summer by metazooplankton ( 0.5 g C m^-2). A highly significant negative correlation was found between protozooplankton and metazooplankton during the plankton growth season. Predation by the latter is thus an important factor regulating size of the protozooplankton population, although other factors also appear to be in operation. Loss rates of the pelagic system through sedimentation are highest in spring and autumn when protozooplankton dominate the grazing community and loss rates are much lower in summer when metazooplankton are the dominant herbivores. Apparently, the impact of protozooplankton grazing on the pelagic system is quite different to that of the metazooplankton.Publication No. 268 of the Joint Research Programme (SFB 95), Kiel University     

Primary production and environmental factors in an oligotrophic biome in the Aegean Sea

The rate of the primary production of the phytoplankton community in the Petalion Gulf, Aegean Sea, was studied from January 1970 to May 1971, at a station situated at approximately Latitude 37°54N; Longitude 24°11E. A variety of physical and chemical parameters such as chlorophyll, primary nutrients (N,P,Si), temperature, salinity, oxygen and light penetration were also studied simultaneously. The rate of the gross primary production varied from 40 to 200 mg C m^-2 day^-1, with a mean value of 90 mg C m^-2 day^-1. The annual gross primary production was calculated to be 33 g C m^-2, which is the minimum known value in the Aegean and Mediterranean Seas. Maximum production was found at the depth of 20 m on the average, mainly due to high light intensities. Petalion Gulf supports a small photosynthetic biomass, as indicated by the low seasonal values of chlorophyll a (0.01 to 0.18 mg m^-2), the highest values being found in the summer. The low production rate noted may have been due to the low nutrient concentrations found: N, 0.04 to 0.32 g-at/1; P, 0.00 to 0.15 g-at/1; Si, 0.45 to 2.25 g-at/1. It is suggested that inorganic phosphorus and nitrogen may alternate in limiting primary production rates in these oligotrophic waters. The temperate waters of the Petalion Gulf are stratified in summer (15.5° to 24.7°C) and well-mixed in winter (12.9° to 15.0°C); they are oxygen-saturated throughout the year, and of high transparency, with 86 m depth for the euphotic zone on the average yearly. The Petalion Gulf is therefore characterized as a typical oligotrophic biome in the Aegean and Eastern Mediterranean Seas.     

Distribution and structure of heterotrophic protist communities in the northeast equatorial Pacific Ocean

The distribution and structure of heterotrophic protist communities and size-fractionated chlorophyll a were studied during the Korea Deep Ocean Study 98 (KODOS 98) research expedition (July 1998) in the northeast equatorial Pacific Ocean (5–11°N). Areas of convergence and divergence formed at the boundaries of the South Equatorial Current (SEC), North Equatorial Current (NEC), and North Equatorial Counter Current (NECC) during the expedition. Water column physicochemical characteristics significantly influenced the size structure of heterotrophic protist communities. Intense vertical mixing and high nutrient and chlorophyll a concentrations characterized SEC and NECC areas, which were affected by converging and diverging water masses, respectively. Nanophytoplankton dominated in SEC and NECC areas; both areas also had relatively high heterotrophic protist biomasses (average 743 µg C m^–2). NEC areas were characterized by a stratified vertical structure, low nutrient and chlorophyll a concentrations, and picophytoplankton dominance. The heterotrophic protist biomass in NEC areas averaged 414 µg C m^–2; nanoprotists (<20 µm) dominated the community. The nanoprotist biomass comprised 49–54% of the total heterotrophic protist biomass in SEC/NECC areas and 67–72% in NEC areas. The biomass of heterotrophic protists was higher in SEC/NECC areas than in NEC areas, but the relative importance of nanoprotists was greater in NEC areas than in SEC/NECC areas. Heterotrophic dinoflagellates were dominant components of the <20 µm and >20 µm size classes in both water columns. The biomass of heterotrophic protists significantly correlated with the net-, nano-, and picophytoplankton biomass in SEC/NECC areas and with the nano- and picophytoplankton biomass in NEC areas. Heterotrophic protists and phytoplankton also showed strong positive correlation in the study area. The size structure of the phytoplankton biomass coincided with that of heterotrophic protists; the heterotrophic protist biomass positively correlated with the protists prey source. These relationships suggest that the community structure of heterotrophic protists and the microbial food web depended on size classes within the phytoplankton biomass. Microzooplankton grazing and phytoplankton growth rates were higher in SEC/NECC areas than in NEC areas. In contrast, the potential primary production grazed by microzooplankton was relatively high in NEC areas (127.3%) compared with SEC/NECC areas (94.6%). Our results indicate that the relative importance and size structure of heterotrophic protists might vary according to two distinct water column structures.Communicated by T. Ikeda, Hakodate