Relationships between heterotrophic bacteria and cyanobacteria in the northern Adriatic in relation to the mucilage phenomenon

High variability of heterotrophic bacterial (HB; 0.1.10(9)-6.10(9) cells L(-1)), nanoflagellates (HNF; 0.02.10(6)-2.4.10(6) cells L(-1)) and cyanobacterial (CB; 10(6)-700.10(6) cells L(-1)) abundances were observed during approximately monthly measurements at six stations along the transect Po Delta-Rovinj from March 1999 to August 2002. Substantially higher values were observed during the stratification period (June-September) in the surface layer of the western stations that were under more direct influence of Po River discharges. Changes of the HB abundance were significantly correlated with temperature in the entire water column and chlorophyll a (Chla) concentration in the surface layer. The nutrients did not look to have directly influenced the HB growth, except orthophosphate in the eastern, more oligotrophic part of transect, where probably HB efficiently competed with phytoplankton for this nutrient. Temperature was also important for CB growth that, however, appeared to occur more intensively in waters with low nutrient concentrations. Probably, in these conditions CB can still develop, while larger autotrophs are strongly nutrient limited. In the upper water column of reduced salinity (35-37), the contributions of CB carbon biomass to the total picoplankton biomass (CB+HB) were mostly larger than 30% (up to 80%) in years with mucilage events (1991, 2000-2002) than in other years. For example, in 1999 only a few values were higher than 30% (up to 50%). In abundant presence of mucilaginous aggregates (e.g. in June 2000 and late June 2002) the chlorophyll a ascribed to CB accounted for much larger portions of total measured Chla. From these results it was concluded that cyanobacteria during mucilage events may play a substantially increased role both as primary producers and prey within the microbial loop of the northern Adriatic.     

Temporal dynamics of dissolved and particulate organic carbon in the northern Adriatic Sea in relation to the mucilage events

The accumulation of dissolved and particulate organic matter may play an important role in mucilage formation in the northern Adriatic. Distributions of dissolved and particulate organic carbon were therefore investigated during the period June 1999-July 2002, when massive mucilage events occurred: in the summer of 2000 and, to a greater extent, of 2002. The seasonal variations in dissolved organic carbon (DOC) concentrations were significant, doubling in summer (up to 150 micromol L(-1)) with respect to winter. The particulate organic carbon (POC) variations were also very large, with a less pronounced seasonal pattern compared to DOC, because the POC changes are much more dependent both on river discharges and on phytoplankton blooms. The comparison of the concentrations between the period before (March-May) and after the onset of mucilage events (June-August) showed that DOC, but particularly POC, were higher in the period before the event of 2002, more markedly in the surface waters of low salinity. The POC increased, reaching mean concentrations of up to 36 micromol L(-1) in March 2002 before the outbreak of the massive mucilage formation in June. This suggests that POC may have a more important role in the mucilage formation than DOC. The highest seasonal variations of organic matter concentrations took place in the upper layer of lower salinity, stressing the importance of stratification and pycnoclines for accumulation and aggregation of the organic matter in the northern Adriatic. The POC contribution to the total organic carbon was low in the oligotrophic waters (DOC/POC ratio >15) and increased with the phytoplankton biomass in the productive waters (DOC/POC ratio <10). Particulate organic carbon predominated over the dissolved inside the mucilage aggregate (DOC/POC ratio <1), probably because aggregation processes, in which colloidal organic carbon is involved, are important. The organic carbon within the aggregates reached a concentration of 13.6 mmol L(-1) which was about 100 times more than in the surrounding waters or in the waters when the mucilages were absent. This indicates that distributions of organic carbon in the northern Adriatic can be extremely patchy during mucilage events.     

Phytoplankton size-distribution and community structure in relation to mucilage occurrence in the northern Adriatic Sea

The spatial and temporal variations of pico-, nano- and microphytoplankton abundance and composition were investigated over a 37 month period, focusing on the ecological role of different size classes of phytoplankton, and on the changes of the community structure that might occur during periods when large mucilage macroaggregates appear. Samples were collected monthly from June 1999 to July 2002 at 11 stations, along three transects covering the northern Adriatic basin. Highest abundances were observed in late-winter/spring for microphytoplankton (mainly diatoms), in spring-summer for nanophytoplankton, and in summer for picophytoplankton. The autotrophic component was more abundant in the summers of 2000 and 2002 (when large mucilage aggregates occurred) than in the summers of 1999 and 2001 (when a massive phenomenon was not observed). This increase was statistically significant for pico-, nano- and, among microphytoplankton, only for dinoflagellates. Blooms of picophytoplankton were often observed at the bottom layer during mucilage summers. The microphytoplankton community during mucilage phenomena was characterized by a species composition (Chaetoceros spp., Cerataulina pelagica, Pseudo-nitzschia delicatissima, P. pseudodelicatissima, Cylindrotheca closterium, Dactyliosolen fragilissimus) comparable to that observed in summers without extensive mucilage occurrence. However, some species appeared with significantly higher densities in the summers of 2000 and 2002: Ceratium furca, C. closterium, Oxytoxum spp., Hemiaulus hauckii and Gonyaulax fragilis. Microscopic observation of aggregates revealed that the microphytoplankton species composition inside the aggregates was comparable to that observed in the water column, with an enrichment of opportunistic species such as C. closterium and P. delicatissima. The presence of mucilage aggregates affects the phytoplankton populations in the water column, even when aggregates are at early stages. It seems that there is a mutual relationship between phytoplankton and aggregates, i.e., several diatom and dinoflagellate species may contribute to the aggregate formation and enlargement, but mucilage aggregates themselves may also affect the phytoplankton populations, allowing the development of a rich diatom community and in general enhancing nanophytoplankton growth.     

Spatial and temporal variability of carbohydrates in the northern Adriatic--a possible link to mucilage events

Large-scale mucilage events in the northern Adriatic have become more frequent during the last few decades. Since carbohydrates (CHO) comprise one of the major constituents of the Adriatic mucilage, studying their seasonal and spatial patterns is one of the key prerequisites for the understanding of this phenomenon. This paper reports on the distribution of particulate (PTCHO) and dissolved carbohydrates (DTCHO) in the period from June 1999 to July 2002, with a special emphasis on the summer mucilage events. A particular attention was paid to the relationship between the phytoplankton crop as reflected by photosynthetic pigments and carbohydrate levels in different seasons. The concentration of PTCHO varied in a very wide range (1-72 micromol C/l). Their seasonal distribution revealed that enhanced levels of PTCHO occurred only in the surface layer (0-5 m) and were associated with major phytoplankton blooms. However, the main pool of CHO was in the dissolved fraction, comprising on average 84+/-10% of the total CHO. In contrast to PTCHO, there was no seasonal coincidence between the enhanced phytoplankton biomass and concentrations of DTCHO. In fact, the concentration of DTCHO showed regularly a marked increase during summer, when the phytoplankton biomass remained relatively low. Nevertheless, the concentration of DTCHO in the productive surface layer, influenced by the Po River discharges, was significantly higher than that in the deeper layers. The detailed vertical distribution of carbohydrates indicated that phytoplankton populations in the bottom layer possessed a rather modest potential as a source of CHO. The spatial distribution in the surface layer along the transect Rovinj-Po River mouth revealed only a moderate increase of the total CHO concentration in the more eutrophic western part of the basin. Analysis of pluriannual variability of the total CHO levels showed that probability of a mucilage event in a given year depended on the accumulation of CHO during spring, but, even more, on the prevailing circulation pattern in the basin during the stratification period.     

Imbalance between phytoplankton production and bacterial carbon demand in relation to mucilage formation in the Northern Adriatic Sea

Spatial and temporal changes in phytoplankton production and bacterial C demand were investigated at four stations in the Northern Adriatic Sea over 3 years. The effect of the Po River plume was observed at the western stations; in particular, the northernmost one (B06) showed the highest values of primary production, both as hourly peaks (up to 14 mg C m(-3) h(-1)) and daily water column integrated values (up to 740 mg C m(-2) day(-1)), the southern station (C04) was only sporadically influenced and did not differ significantly from the easternmost ones (C12 and B13), where the lowest TPP values were recorded (around 1 mg C m(-3) h(-1)). In this study the first in situ data are reported on short-term phytoplankton C extra cellular release in the Northern Adriatic Sea. At every station a considerable percentage of primary production (PER>20% as an average, with peaks of up to 70%) was released as dissolved organic carbon. In particular, an association of fairly high PER (>10%) and specific production (Pb>10 mg C mg chl(-1) h(-1)) was observed from spring to summer, when the mucilage phenomenon usually starts. This result might suggest the presence of an uncoupling between photosynthesis and growth, probably related with nutrient availability, which would be responsible for a high production of extra cellular organic carbon. Phytoplankton primary production and bacterial carbon production were closely related and bacterial C production accounted, on average, for a higher percentage of primary production than the values typically reported in the literature on aquatic environments. The flow of organic matter from phytoplankton to bacteria seems to satisfy the bacterial carbon demand during most of the spring and summer, at least in the upper water layers. However, during the summer, there is evidence that BCD sometimes exceeds the amount of C produced by phytoplankton. Neither phytoplankton nor bacterial production showed significant differences over the relevant years, and their absolute values did not change when comparing periods with or without mucilage. However, there were indications of an uncoupling between phytoplankton photosynthesis and growth and of a shift from an autotrophic to a heterotrophic metabolism, especially during the spring and summer period when mucilage might occur.     

Meteorological and oceanographic conditions in the northern Adriatic Sea during the period June 1999-July 2002: influence on the mucilage phenomenon

Mucilage events (formation of very large organic aggregates and gelatinous surface layers) have been documented several times during the past two centuries in the northern Adriatic Sea (NA), while their frequency has significantly increased since 1988. In this work, meteorological and oceanographic conditions in the NA during the period June 1999-July 2002 are described and their relation to the outbreak and fate of the mucilage phenomenon was investigated. Salinity and temperature data were collected during approximately monthly cruises along three transects in the NA. Relevant meteorological situations (air temperature, rainfall, wind) were selected from large-scale ECMWF analyses and from the Local Analysis and Prediction System (LAPS; Emilia Romagna Meteorological Service), while sea conditions (waves) were analysed by means of the Wave Adriatic Model (WAM). Data for air temperature, rainfall, and wind from several meteorological stations in the region were used. Average seasonal cycles of sea temperature and salinity simulated with statistical models, based on historical data collected in the NA since 1972, were used to determine thermal and haline anomalies. The monthly anomaly variability of maximum and minimum air temperatures, rainfall amount and number of rainy days did not appear to be relevant for the mucilage phenomenon outbreak. In contrast, both vertical and horizontal thermohaline gradients in the region were more developed during late spring and summer of 2000 and particularly of 2002, when the mucilage events were of greatest extent in space and time, compared to 2001 (short-lived event) and 1999 (no event). These more pronounced gradients were due to a combination of several unusual conditions: sharp heating of the sea surface in May-June, domination of eastwards transport of freshened waters formed in the Po Delta area, and intrusion of very high salinity intermediate waters originating in the eastern Mediterranean. Moreover, in winter of both 2000 and 2002 very dense and cold water formed and remained in the bottom layer until spring, contributing to increase the stratification degree of the water column. The duration of the mucilage events and their spatial distribution in the region depend strongly on meteorological changes. Recurrent anticyclonic conditions, characterized by low wind and calm sea, favour extended events in time (up 2 months in 2002). In contrast, highly perturbed weather, particularly due to strong "bora" wind, can be determined in sharp decay of the event (e.g. in July 2000).     

Relevance of the dinoflagellate Gonyaulax fragilis in mucilage formations of the Adriatic Sea

Oceanographic cruises were carried out monthly from June 1999 to July 2002 to follow the mucilage formation process in the Northern Adriatic Sea. Results show that in correspondence with these events the dinoflagellate Gonyaulax fragilis (Schütt) Kofoid was observed both in the water column and within mucilage aggregates. In the water column, increasing abundances were observed from May until July, with values never exceeding 8500 cells l(-1). Much higher densities were observed within superficial gelatinous aggregates (22800-3400000 cells l(-1)). In mucilage samples, a large number of decomposing cells were present, together with abundant alive cells, enveloped in exudates. G. fragilis isolated from mucilage samples was cultured in three different culture media; it was characterized by a low growth rate but it produced a high amount of polysaccharides. The highest yield both in terms of cell number and carbohydrate production was observed in the medium having the highest nitrogen and phosphorus content and the lowest N/P ratio. The monomeric composition of G. fragilis carbohydrates, compared with that of mucilage samples, showed that in both natural and cultured samples galactose was the most abundant sugar; in addition, an overall good correlation, especially between the monomeric carbohydrate composition of G. fragilis grown in f/2 medium and that of a mucilage sample in which this species was present in high density, was observed.     

Exo-enzymatic activities and dissolved organic pools in relation with mucilage development in the Northern Adriatic Sea

We tested the hypothesis that the appearance of mucilage in the Northern Adriatic Sea was related with the accumulation of dissolved organic compounds released by intensive enzymatic activities and not utilized as direct substrate for microbial growth. To do this enzymatic activities and dissolved organic and inorganic pools in periods characterized by the presence of mucilage and in the same seasons but in absence of mucilage were compared. Extracellular enzymatic activities (aminopeptidase, beta-glucosidase and alkaline phosphatase), nutrient pool concentrations (total dissolved nitrogen, dissolved organic nitrogen, total dissolved phosphorus, dissolved organic phosphorus) and the biochemical composition of particulate and dissolved organic matter (in terms of proteins and carbohydrates) were determined on a monthly basis over a period of 3 years. Aminopeptidase and alkaline phosphatase activities displayed higher values in springs preceding the appearance of mucilage than in spring when no mucilage was observed. Beta-Glucosidase activity showed significantly higher values in summer periods characterized by the massive production of mucilage than in summers without mucilage events. The months preceding mucilage events were also characterized by an increase of the alkaline phosphatase to aminopeptidase activity ratio and by a significant accumulation of dissolved proteins. These findings, together with the significant increase of the DON/DOP ratio, suggest that mucilage formation is favoured by the deficiency of organic P. The present study provides compelling evidences that mucilage formation is favoured by the unbalance between organic matter mobilization by enzymatic activities and the accumulation of labile dissolved organic-N compounds.     

Inter-annual variations of planktonic food webs in the northern Adriatic Sea

The temporal dynamics of microphytoplankton, microzooplankton and mesozooplankton were monitored over 37 months in the Adriatic Sea in order to identify alterations in the plankton structures, which can lead to, or enhance the production of macro-aggregates, that affected the entire northern basin in summers 2000 and 2002, and to assess any negative effects of mucilage on plankton temporal patterns. Samples were collected monthly, from June 1999 to July 2002, on three transects at 9 stations across the northern and central Adriatic Sea. Besides the high year-to-year variations in abundances and taxonomical composition, plankton communities only showed a clear seasonal succession during 2001, when since April a grazing food web developed and was able to control large sized phytoplankton increase. In spring-summer 2000 and 2002 consumer abundances remained quite low and the dominant mesozooplankton summer species (Penilia avirostris) did not reach its usual summer maximum. The lack of an efficient top control was more evident on the northernmost transect, where generally grazing food web prevails over the microbial one. A large part of the microphytoplankton blooms, although not particularly intense, was exported to the bottom in the particulate phase, where it was processed by bacteria, enhancing the production of refractory dissolved material.     

Mercury and methylmercury in the Gulf of Trieste (northern Adriatic Sea)

The distribution, sources and fate of mercury (Hg) in the water column of the Gulf of Trieste (northern Adriatic Sea), affected by the Hg polluted river Soca/Isonzo for centuries draining the cinnabar-rich deposits of the Idrija mining district (NW Slovenia), were studied in terms of total and dissolved Hg, reactive Hg, total and dissolved methylmercury (MeHg), mesozooplankton Hg and MeHg, and sedimentation rates of particulate Hg. Higher total Hg concentrations in the surface layer were restricted to the area of the Gulf in front of the river plume expanding in a westerly direction. Higher concentrations in bottom water layers were the consequence of sediment resuspension. Dissolved Hg exhibited higher concentrations in the surface layer in the area in front of the river plume. Higher bottom concentrations of dissolved Hg observed at some stations were probably due to remobilization from sediments, including resuspension and benthic recycling. The relationship between dissolved Hg in the surface layer and salinity showed nonconservative mixing in June 1995 during higher riverine inflow and nearly conservative mixing in September 1995 during lower riverine inflow. Both mixing curves confirm the river Soca/Isonzo to be the most important source of total and dissolved Hg, which are significantly correlated, in the Gulf. Reactive Hg is significantly correlated with dissolved Hg, indicating that the majority of dissolved Hg is reactive and potentially involved in biogeochemical transformations. The higher total MeHg in the bottom layer is the result of remobilization of MeHg from sediments including benthic fluxes. Strong seasonal variation of sedimentation rates of particulate Hg was found during a 2-year study in the central part of the Gulf. These variations followed those of total sedimented matter, indicating that sedimented Hg is mostly associated with inorganic matter. About a 2.5-fold higher fluxes of particulate Hg were observed at the depth of 20 m relative to 10 m which is attributed to bottom sediment resuspension. Temporal variability of mesozooplankton Hg and MeHg is the consequence of biomass and species variations, and grazing behaviour. From the preliminary Hg mass balance it appears that the Gulf is an efficient trap for total Hg and a net source of MeHg.     

Chemical and biochemical parameters of cultured diatoms and bacteria from the Adriatic Sea as possible biomarkers of mucilage production

Bacteria and diatom strains from the Adriatic Sea were investigated, under standard and altered environmental conditions, for carbohydrate production and for the presence of specific biomarkers. Algae from P-depleted cultures showed an increase in extracellular carbohydrate production, a significantly lower chlorophyll a content and unchanged total lipid levels. However, the fatty acid composition of algal cultures was severely affected by low P levels, in that, total saturated and monounsaturated fatty acids increased and total polyunsaturated fatty acids decreased. Marine heterotrophic bacteria resulted enriched by 4 to 6 orders of magnitude in mucilage samples respect to surrounding seawater, unlike other groups of bacteria such as the non-halophylic heterotrophs. The major fatty acids detected in bacteria were 16:0 and 18:1n-7; the uneven fatty acids 17:0i, 17:0 and 17:1 also constituted an important component of various strains and, as a result, the total monounsaturated fraction represented the main component of total fatty acids. All the mucilage samples analysed shared the same general fatty acid composition features with a high amount of saturated components, especially 16:0; typical marine polyunsaturated fatty acids, such as 20:5n-3 and 22:6n-3, were found at very low levels. With regard to the sterol composition, the analysed algal species and bacteria showed that different compounds prevailed in the different species, and under P-deprivation sterol distribution resulted differently affected in the various algal species. In mucilage samples an overall prevalence of cholesterol was observed and, among 4alpha-methylsterols, constantly present, dinosterol prevailed in all samples. Vibrational IR spectroscopic analyses confirmed the main results obtained with the GC analysis: a higher unsaturation degree in nutrient replete diatom cultures than in P-depleted ones, a lower amount of P-containing compounds in the latter, bacterial lipid profiles with a high amount of free carboxylic acids and/or ketones and a low unsaturation degree and, finally, mucilage samples with a very low unsaturation degree. All these results allowed some speculations on the involvement of the various microbial and phytoplankton components in mucilage genesis.     

Extracellular enzyme activity and dynamics of bacterial community in mucilaginous aggregates of the northern Adriatic Sea

Bacterial degradation of mucilaginous aggregates (creamy layers, stringers and macroflocs) collected during two summer events (2001-2002) was tested. The objective was to describe the temporal trend of the bacterial activity, abundance and composition in the aggregated and dissolved organic matter under different trophic conditions. In the native aggregates proteins and organic phosphorous were actively hydrolyzed as aminopeptidase and alkaline phosphatase activities represented up to 87% and 25% of total activity, respectively; polysaccharides were less hydrolyzed and the highest activities were observed for beta-glucosidase (5% of the total). This hydrolysation pattern tends to a progressive accumulation of long persistent polysaccharides. During short term incubations nutrient addition (P, N and Glucose) differently stimulated bacterial growth in the seawater: P played the main role in stimulating bacterial production from 3 to 6 folds higher than in the control, whereas a secondary C-limitation was observed only for bacteria growing on seawater from macroflocs. This scarce dissolved organic carbon (DOC) bioavailability was confirmed by the lower DOC removal (13% macroflocs, 36% stringers). The total amount of carbon incorporated by bacteria living on aggregates was similar (0.58 mg C L(-1)) both in the control and under P enrichments showing a more balanced condition with respect to the seawater. Hence the well-known P limitation in the Northern Adriatic Sea affects only dissolved organic carbon uptake without influencing the uptake of aggregated organic matter. Organic matter limitation was observed only on stringers--total C incorporated raised to 0.96 mg C L(-1) after PNG addition. Macroflocs release of refractory compounds leads to DOC accumulation (73 microM DOC) contributing to inflate the pool of refractory DOC in the surrounding waters. Several evidences, including different monosaccharide composition of stringers and macroflocs (glucose 15% and 56% on stringers and macroflocs, respectively), bring to the conclusion that stringers are in an older stage in comparison with macroflocs. Community composition described by fluorescence in situ hybridization did not show significant differences between free-living and attached bacteria but it was modified by the different enrichment conditions: Cytophaga-Flavobacteria increased after inorganic nutrients enrichments while organics advantaged gamma-Proteobacteria.     

A numerical study of the interannual variability of the Adriatic Sea (2000-2002)

A free-surface, three-dimensional finite-difference numerical model based on the Princeton Ocean Model (POM) has been implemented in order to simulate the interannual variability of the Adriatic Sea circulation. The implementation makes use of an interactive surface momentum and heat flux computation that utilizes the European Centre for Medium-Range Weather Forecasts (ECMWF) 6-h analyses and the model predicted sea surface temperatures. The model is also nested at its open boundary with a coarse-resolution Mediterranean general circulation model, utilizing the same surface forcing functions. The simulation and analysis period spans 3 years (1 Jan 2000 to 31 Dec 2002) coinciding with the "Mucilage in the Adriatic and the Tyrrhenian" (MAT) Project monitoring activities. Model results for the simulated years show a strong interannual variability of the basin averaged proprieties and circulation patterns, linked to the atmospheric forcing variability and the Po river runoff. In particular, the years 2000 and 2002 are characterized by a weak surface cooling (with respect to the climatological value) and well-marked spring and autumn river runoff maxima. Conversely, 2001 is characterized by stronger wind and heat (autumn cooling) forcings but no river runoff autumn peak, even though the total amount of water inflow during winter and spring is sustained. The circulation is characterized by similar patterns in 2000 and 2002 but very different structures in 2001. During the latter, deep water is not formed in the northern Adriatic. A comparison with the observed data shows that the major model deficiencies are connected to the low salinity of the waters, probably connected to the missed inflow of salty Ionian waters of Aegean origin and to the numerical overestimation of the vertical mixing processes.     

Interannual variability of surface heat fluxes in the Adriatic Sea in the period 1998-2001 and comparison with observations

Surface heat fluxes of the Adriatic Sea are estimated for the period 1998-2001 through bulk formulae with the goal to assess the uncertainties related to their estimations and to describe their interannual variability. In addition a comparison to observations is conducted. We computed the components of the sea surface heat budget by using two different operational meteorological data sets as inputs: the ECMWF operational analysis and the regional limited area model LAMBO operational forecast. Both results are consistent with previous long-term climatology and short-term analyses present in the literature. In both cases we obtained that the Adriatic Sea loses 26 W/m2 on average, that is consistent with the assessments found in the literature. Then we conducted a comparison with observations of the radiative components of the heat budget collected on offshore platforms and one coastal station. In the case of shortwave radiation, results show a little overestimation on the annual basis. Values obtained in this case are 172 W/m2 when using ECMWF data and 169 W/m2 when using LAMBO data. The use of either Schiano's or Gilman's and Garrett's corrections help to get even closer values. More difficult is to assess the comparison in the case of longwave radiation, with relative errors of an order of 10-20%.     

Behaviour of Hg species in a microtidal deltaic system: the Isonzo River mouth (northern Adriatic Sea)

Mercury species in the highly stratified water column of the Isonzo River mouth (northern Adriatic Sea) were investigated in February, May and August 2002 (low-normal fluvial discharge). Total, dissolved and particulate mercury (Hg) and methylmercury (MeHg) were measured and their concentrations were related to physico-chemical parameters in terms of temperature, salinity, turbidity, O(2) concentrations and total suspended matter (TSM) as well as particulate organic carbon (POC) content. Particulate Hg and MeHg are well correlated to medium-fine silty suspended sediment and organic matter, respectively. Desorption of Hg from particles in the brackish layer was observed. Due to the presence of a saltwedge, the lower river course seems to be a trap for Hg carried by fluvial waters. MeHg normally decreases approaching the sea but the local increase in the inner zone of the saltwedge could be related to methylation processes in the bottom water layer. Hg export is effective only during medium and high riverine flows acting as an important source of Hg into the northern Adriatic and the adjacent lagoon system.     

Circulation and horizontal fluxes in the northern Adriatic Sea in the period June 1999-July 2002. Part II: nutrients transport

Nutrient concentrations (orthophosphate, orthosilicate and dissolved inorganic nitrogen-DIN) were measured on three transects in the northern Adriatic approximately monthly in the period June 1999-July 2002. The net nutrient transports across the three transects were estimated from these concentration values and calculated water fluxes [Grilli F, Paschini E, Russo A, Precali R, Supi? N. Circulation and horizontal fluxes in the Northern Adriatic Sea in the period June 1999_July 2002. Part I: geostrophic circulation and current measurement. Sci Total Environ 2005-this issue], with the aim to verify their possible role in the mucilage phenomenon. The nutrient transports in the northern Adriatic were very variable, seasonally and from year to year, both in intensity and direction. Some differences were noticed also among the nutrient species. At the northernmost transect Po Delta-Rovinj minimal values and variable directions were recorded, probably due to the prevailing of eastward transversal transports, observed already in late winter, and generally dominant in spring and summer. Northward transport was often measured at the central transect Cesenatico-Cape Kamenjak, particularly in spring 2000 and 2002, and summer 2001, as well as for only DIN in summer 1999. In contrast, southward directions prevailed at the southernmost transect Senigallia-Susak Island. In October significant southward transports occurred at all transects and for all nutrients, particularly strong in 2000, when exceptionally high Po River discharges occurred, and the Western Adriatic Current (WAC) was well developed. However, the nutrient transports in the upper water column were low or northwards at both transect Cesenatico-Cape Kamenjak and Senigallia-Susak Island in June and July 2000, 2001, and 2002, when the mucilage phenomenon developed, but were opposite in 1999, a year without events. The results suggest that high variations of nutrient fluxes and their ratios, in conditions of reduced water dynamics, are essential for the development of the phenomenon, rather than the absolute amounts of the nutrient inputs.     

Circulation and horizontal fluxes in the northern Adriatic Sea in the period June 1999-July 2002. Part I: geostrophic circulation and current measurement

The dramatic increase in the occurrence of massive mucilage events in the northern Adriatic (NA) since their recent conspicuous reappearance in the late 1980s prompted a study of circulation and horizontal fluxes. Three transects with equidistant stations (10 km) were thus monitored monthly between June 1999 and July 2002. The geostrophic method was used to compute currents across the three transects from the CTD data, and dynamic heights provided a picture of the horizontal surface circulation. Currentmeter data records were used to adjust the reference surface and to validate the results for the southernmost and deeper (up to 70 m) transect (Senigallia-Susak Island). Geostrophic currents allowed estimation of monthly water fluxes across the transect. Different circulation regimes in the NA were observed, which may have affected mucilage events. When mucilage was absent (1999) or reduced (2001) in the western sector, the Western Adriatic Current (WAC, carrying water out of the NA) was found to be active, whilst the WAC was very weak or reversed when massive mucilage events occurred (2000 and 2002). Opposite behaviour has been observed for the Istrian Coastal Counter-Current (ICCC, retaining freshwater water in the NA) which was more intense during or after massive mucilage events and did not appear when mucilage was absent. Both WAC weakening and ICCC strengthening indicate a longer residence time of riverine waters in the NA, which favours mucilage development. Conclusively, WAC and ICCC result as key elements in controlling massive mucilage phenomena in the NA.     

Changes in nutrient concentrations and ratios during mucilage events in the period 1999-2002

Nutrient and chlorophyll a concentrations and salinity were measured, approximately monthly, from June 1999 to July 2002 at 20 stations along three transects in the northern Adriatic Sea, north of the line Susak Island-Senigallia, with the aim of confirming or rejecting the hypothesis that changes in nutrient ratios may have an important role in the mucilage phenomenon. The data analyses were focused on the two major water types identified in the region: lower salinity (32-37) and oxygenated surface waters (type 1) in which the mucilage phenomenon primarily developed, and high salinity water originating from other parts of the Adriatic (type 4). Marked variability of dissolved inorganic nitrogen (DIN) in type 1 waters was roughly correlated with extreme fluctuations of the Po River flow rate during the investigated period. In contrast, the orthophosphate (PO4) concentration was primarily controlled by phytoplankton assimilation. The nutrient discharges and DIN/PO4 ratios (median 120) in the freshened surface layer were much higher and more variable in the period before the mucilage event in 2001 than in 2000 (median 75), and particularly in 2002 (median 30), although intensity and duration of the 2001 event were the lowest. However, in that period of 2000 and 2002 significant transversal transport of freshened waters occurred, despite the unusually low Po flow rates. In summer, in conditions of low freshwater discharge and the prevailing of semi-enclosed circulation in the region, more efficient DIN assimilation by phytoplankton occurred, probably due to a faster recycling of PO4. However, in 2002 this process appeared to have already started in March. Changes of the orthosilicate (SiO4)/DIN ratio were mainly dependent on DIN concentrations. In the more saline waters (type 4) the nutrient concentrations, particularly DIN, were much lower and no significant relationships were noticed among the studied parameters. Nutrient concentration and ratio changes do not trigger mucilage events, although very probably they have an essential role in combination with several other physical (pulsing freshwater discharge, marked stratification, minimal water exchange) and biological (e.g., increased plankton excretion, limited bacterial degradation) factors.     

Benthic fluxes of cadmium, lead, copper and nitrogen species in the northern Adriatic Sea in front of the River Po outflow, Italy

Trace heavy metal (Cd, Pb and Cu) and nitrogen species (N-NO3, N-NO2 and N-NH4) fluxes between sediment and water were examined for approximately 4 days, in a coastal marine station located in the northern Adriatic Sea in front of the River Po outflow. An in situ benthic chamber, equipped with electronic devices for monitoring and adjustment of oxygen and pH and with a temperature detector, was used. The benthic chamber experiment enabled study of the temporal trend of metals and nutrients when oxygen concentration varied in a controlled environment. Although particular care was devoted to chamber deposition and parameter control, sediment resuspension occurred at the beginning of the experiment and O2 fluctuations were observed during the course of the experiment. Pb concentration was affected by both resuspension and oxic conditions in bottom water, which prevented determination of any reasonable Pb flux value. Cd and Cu, not influenced by oxygen fluctuations, reached an equilibrium phase in a short period with initial positive fluxes from sediment of 0.68 (S.D. = 0.07) and 6.9 (S.D. = 5.6) pmol cm(-2) h(-1), respectively. With regard to nitrogen species, the highest positive flux was that of N-NH4 (10.5, S.D. = 2.4, nmol cm(-2) h(-1)) whose concentration increased in the chamber, while nitrate concentration (initial flux of -5.7, S.D. = 1.5, nmol cm(-2) h(-1)) immediately decreased after the beginning of the experiment. Nitrite concentration was almost constant throughout the experiment and its flux was generally low (initial flux 0.1, S.D. = 0.9, nmol cm(-2) h(-1)).