Limiting nutrients of phytoplankton community in Hiroshima Bay,Japan

Ratios of DIN:DIP, indigenous algal assay and nitrogen fixation by phytoplankton were determined to estimate the limiting nutrient for phytoplankton growth in Hiroshima Bay, Seto Inland Sea, Japan. The algal assay was performed using indigenous pico-, nano- and microphytoplankton assemblages fractionated by filtration. Nitrogen fixation was determined using an acetylene reduction method. The DIN:DIP ratio in the study area was above 16:1 (Redfield ratio) in August 1993 and May 1994. However, in other months, the ratio was below 16:1. These values suggest that the growth of phytoplankton community is limited by nitrogen, with an exceptional phosphorus limitation in August 1993 and May 1994. Nutrient enrichment algal assay of size-fractionated phytoplankton community also supported these findings. The growth of picophytoplankton fraction was stimulated by phosphorus addition in August 1993 and May 1994, while nano- and microphytoplankton fractions were stimulated by nitrogen addition in almost all months during the study period. No nitrogen fixation by phytoplankton was detected during the study period. The results suggest that nitrogen rather than phosphorus is the limiting nutrient for the growth of the whole phytoplankton community, but picophytoplankton by itself, tends to be limited by phosphorus compared to other large-size phytoplankton when high DIN:DIP ratio was observed.     

Fatty acid profiling of microbial community during aging of mucilaginous aggregates in the northern Adriatic

Aggregates differing in size and phytoplankton community composition were sampled in winter/spring 2001 and summer 1997 and 2002 (during mucilaginous event) in the northern Adriatic Sea. The fatty acid profiles (FAME) were determined in aggregates, bacterial population was grown from each aggregate and each bacterial isolate from aggregate's plated cultures. All aggregates irrespective of the season, aggregate size or phytoplankton community composition contained isolates from three distinct groups, with fatty acid profiles corresponding to alpha-Proteobacteria, gamma-Proteobacteria and Cytophaga-Flavobacter (CF) complex but in different relative proportions. Fatty acid profiles of the bacterial population grown from each aggregate revealed either the domination of one group (gamma, alpha or CF) or contribution of more groups (gamma and CF; alpha and CF). Specifically, as the aggregates age a shift in favor of bacteria belonging to CF-complex and corresponding decrease in bacteria similar to that of Proteobacteria occur. During the aggregate aging process and degradation of the mucous matrix, besides bacterial succession, the phytoplankton growth took place inside the aggregates. Such an aging pattern was confirmed through laboratory experiments with aggregates inhabited by the diatom Cylindrotheca closterium.     

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.     

Mucilage microcosms

In the summers of 2000, 2001 and 2002, large amounts of sticky mucilaginous material aggregated to form masses of impressive dimensions over large areas of the Adriatic Sea, particularly in its northern part. Aggregates differing in size were sampled by SCUBA divers and submitted to chemical (nutrient and organic matter concentrations) and biological analysis (virus, bacteria and phytoplankton abundances and bacterial metabolism). Suspended and sinking mucilaginous aggregates were biota-rich environments where the abundance of planktonic organisms and the concentration of nutrients were orders of magnitude higher than in the surrounding seawater. The embedded phytoplankton was mostly composed of diatoms, but the dinoflagellate Gonyaulax fragilis, previously reported in association with the aggregates, was also present. A variety of processes occurred within the mucilaginous aggregates which resulted in the transformation of the organic matter composition and lability and contributed to a partial degradation of mucilage. For an efficient degradation of mucilage, several conditions are necessary: high bacterial abundance and activity and an efficient recycling of nutrients within the aggregates. Most of these conditions, appear to change depending on the type and age of the aggregate. During the first phase of aggregation (cobwebs and ribbons), bacterial activities addressed the degradation of organic matter, particularly that of the nitrogen fraction. The degradation products were rapidly taken up by bacteria, supporting an increase in their abundance and production. In aged mucilage (clouds), the degradation processes decreased and the bacterial metabolism suggested the presence of new organic labile compounds probably due to phytoplankton production. On the basis of our results, stringers, generally considered the first step of the aggregation process, seemed to be the result of a mechanical disruption of other types of aggregates.     

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.     

Macrobenthos community structural changes off Cesenatico coast (Emilia Romagna, Northern Adriatic), a six-year monitoring programme

Soft bottom macrobenthos at a station located off Cesenatico (Emilia Romagna, Northern Adriatic Sea) was investigated seasonally for six years from July 1996 to July 2002. Species composition and abundance of the community have been studied in relation to fluctuation in the water environment parameters, sediment texture patterns and mucilage, that occurred mainly in the water column at the study site. Three major Po river flow peaks occurred in November 1996, October 2000 and May 2002; after these events the community was reduced to minimum abundance values (total density<2000 individuals m(-2)). In the period between the first two episodes the river discharge remained rather low and conditions of increased salinity, lower nutrients and chl a and good oxygen saturation were experienced. The fossorial Crustacean Ampelisca diadema became dominant in the community between the first two river flow events, reaching maximum density of 10,200 individuals m(-2) and substituting the bivalve Corbula gibba, indicator of sediment instability. Species richness increased in the same period. The role of Ampelisca as a facilitator in structuring the community is discussed. Corbula gibba never recovered to initial densities, apart from an abundance peak that occurred in the summer of 2000. Faunal composition seemed to evolve slowly towards a higher degree of structural complexity (positive trend in diversity and evenness index). In the study site near-bottom mucilage events occurred in the summers of 1997, 1998, 2002; they appeared uncorrelated with the observed changes in the community structure. Multivariate analysis of community structure (MDS, ANOSIM) illustrates that community changes in this station are driven mainly by hydrographical conditions influencing sediment texture patterns and trophic resources for the benthos.     

Transparent exopolymeric particles' distribution in the northern Adriatic and their relation to microphytoplankton biomass and composition

Transparent exopolymer particles (TEP) are generated during transformation processes within the non-living organic matter continuum in the seawater. Seasonal dynamics of TEP along the Po River Delta-Rovinj transect was followed during the 3-year period, using a spectrophotometric method of determination of TEP. Relationships between TEP and phytoplankton biomass (chlorophyll a), abundance and composition, as well as phaeophytin, were analysed. Microphytoplankton spring blooms are confirmed to be an important source of TEP, with some species more related to TEP concentration dynamics, such as Chaetoceros sp. and Skeletonema costatum. Notable spatial west to east gradient of autotrophic biomass and consequently of TEP was observed, as well as gradual decrease with depth. In addition, TEP concentration increments were observed up to 2 months before the mucilage events followed by the decrease of TEP concentration during the event itself. Pattern of TEP dynamics in the years with mucilage events was the most pronounced in 2002, when the event was spatially and temporally the most extended.     

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.     

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.     

Mucilaginous aggregates in the northern Adriatic in the period 1999-2002: typology and distribution

The spatial and temporal distributions of different types of visible mucilaginous aggregates were investigated monthly by underwater video cameras in the northern Adriatic (NA) from June 1999 to July 2002. Small aggregates (flocs, macroflocs and stringers) were observed in all seasons with higher values in autumn and winter. Larger aggregates (ribbons, cobwebs, clouds, and false bottoms) formed only during late-spring and summer, particularly in 2000 and 2002, when the seasonal thermohaline stratification was marked. In fact, the process of mucilage formation took place mainly in the upper water column, above the main pycnocline. The spatial distribution shows that higher concentrations of small aggregates occur in the more productive coastal waters. The larger aggregates form or accumulate mainly in coastal waters in the southern part of the NA and in the central part of the Po River delta-Rovinj transect, where a gyre greatly influences the aggregates distribution. The fronts between low salinity coastal waters and high salinity waters of southern origin play an important role for accumulation and aggregation of the mucilage during spring and summer. Most of the aggregates accumulate in correspondence with strong pycnoclines with differences in density anomaly of 2 kg m(-3) or higher. False bottoms formed in correspondence with strong haloclines (DeltaS congruent with 2), while cobwebs and ribbons also occurred when the halocline was less marked (DeltaS<0.5). Meteorological conditions greatly influenced the aggregate formation. Calm weather and weak winds favour aggregation processes, while strong north-easterly winds, causing intense mixing, result in the dispersion of the mucilaginous aggregates, as occurred during the Bora event in July 2000.     

Chemical characterization of different typologies of mucilaginous aggregates in the Northern Adriatic Sea

The chemical composition of mucilage aggregates found during summer 2000, 2001 and 2002 in the North Adriatic Sea depends on the nature of the organic matter during aggregation, on the environmental conditions of the site of formation and on the transformations during ageing. The mucilages were composed of organic matter, together with a significant inorganic fraction. Elemental analysis revealed 12.5-32.2% of organic carbon, 0-7.3% of inorganic carbon and 1.0-3.7% of nitrogen. The C(org)/N ratios of most aggregates were between 7.5 and 12.6, values close to those found in the suspended matter; higher ratios were found in large-size (>5 m) aggregates which are probably older. The content of carbohydrates and proteins determined in the aggregates, respectively, 15.4+/-8.9% and 7.9+/-4.8%, w/w, showed a prevalence of carbohydrates over proteins. Neutral carbohydrate analysis of purified polysaccharides from mucilage samples showed very similar signatures with high relative abundance of galactose and glucose. Humic, fulvic and humin substances extracted from the mucilages constitute an important fraction of the organic matter in the aggregates. The humin (a fraction insoluble in acidic and basic media) was present in all mucilage samples, indicating the refractory nature of a part of the organic matter in the mucilage. The iron and calcium could play a role during the aggregation process to form a complex with polysaccharides and humic fractions. The C(org)/N ratio 10+/-2 found in the humic acids extracted from the Adriatic aggregates disclosed a marine origin. The low phosphorus content and the high C(org)/P ratio found in the aggregates might depend from high bacteria activity or from the aggregation of organic fractions depleted of phosphorus. The principal inorganic species contained aluminium and silicon, part of which was of biogenic origin and was more significant in the offshore mucilage aggregates than in the coastal ones. The Si(biog)/C(org) ratio showed that diatoms were always present in the aggregates, although it cannot be established whether these are the producers or these develop within the aggregates.     

Phytoplankton community succession shaping bacterioplankton community composition in Lake Taihu, China

PCR-denaturing gradient gel electrophoresis (DGGE) and canonical correspondence analysis (CCA) were used to explore the relationship between succession of phytoplankton community and temporal variation of bacterioplankton community composition (BCC) in the eutrophic Lake Taihu. Serious Microcystis bloom was observed in July-December 2008 and Bacillariophyta and Cryptophyta dominated in January-June 2009. BCC was characterized by DGGE of 16S rRNA gene with subsequent sequencing. The DGGE banding patterns revealed a remarkable seasonality which was closely related to phytoplankton community succession. Variation trend of Shannon-Wiener diversity index in bacterioplankton community was similar to that of phytoplankton community. CCA revealed that temperature and phytoplankton played key roles in structuring BCC. Sequencing of DGGE bands suggested that the majority of the sequences were affiliated with common phylogenetic groups in freshwater: Alphaproteobacteria, Betaproteobacteria, Bacteroidetes and Actinobacteria. The cluster STA2-30 (affiliated with Actinobacteria) was found almost across the sampling time at the two study sites. We observed that the family Flavobacteriaceae (affiliated with Bacteroidetes) tightly coupled to diatom bloom and the cluster ML-5-51.2 (affiliated with Actinobacteria) dominated the bacterioplankton communities during Microcystis bloom. These results were quite similar at the two sampling sites, indicating that BCC changes were not random but with fixed pattern. Our study showed insights into relationships between phytoplankton and bacterioplankton communities at species level, facilitating a better understanding of microbial loop and ecosystem functioning in the lake.     

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.     

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.     

Mucilage impact on gorgonians in the Tyrrhenian sea

The mucilage phenomenon has affected the Tuscan Archipelago since its first appearance (1991) in the Tyrrhenian Sea (Mediterranean Sea) [Innamorati M, Raddi E, Buzzichelli A, Melley S, Demoulin M. Le mucillaggini nel Mar Tirreno. Biol Mar Suppl Notiz 1992;1:23-26; Sartoni G, Sonni C. Tribonema marinum J. Feldmann e Acinetospora crinita (Carmichael) Sauvageau nelle formazioni mucillaginose bentoniche osservate sulle coste toscane nell'estate 1991. Inf Bot Ital 1991;23:23-30; Sartoni G, Cinelli F, Boddi S. Ruolo di Tribonema marinum J. Feldmann ed Acinetospora crinita (Carmichael) Sauvageau negli aggregati mucillaginosi bentonici delle coste toscane. Biol Mar Suppl Notiz 1993;1:31-34]. Seasonally, these mucous aggregates become very common in the benthic domain. The gorgonians are the most exposed organisms because they provide the best support for mucilage growth; indeed, their long branches positioned in orthogonally with respect to the current so as to capture plankton, also trap the filamentous mucilage present in the water. A 3-year monitoring programme at Capo Calvo (Island of Elba) was carried out in order to study the appearance of the mucilage phenomenon and its impact on three species of gorgonians (Eunicella cavolinii, Eunicella singularis, and Paramuricea clavata). The composition of mucilage and the gorgonian recovery capacity, when damaged, were investigated. During the first year of study (1999), no relevant interactions between gorgonians and mucilages were recorded. Instead, massive presence of mucilages causing different types of damage to the different gorgonian species investigated was recorded during the second (2000) and third year (2001). The type and the extent of the impact of mucilages also depend on the season. Three species of algae (Nematochrysopsis marina, Chrysonephos lewisii and Acinetospora crinita) constitute the principal components of the mucilaginous aggregates. In general, the first two species occur during the spring season, down to 20 m, while A. crinita occurs at greater depths. In July, when the mucilages reach their maximum development, C. lewisii is the predominant species. This species mainly affects E. cavolinii and E. singularis while A. crinita mainly affects P. clavata, which colonizes greater depths.     

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.     

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.     

Relationships between northern Adriatic Sea mucilage events and climate variability

A long term analysis (1865-2002) of meteorological data collected in the Po Valley and Northern Adriatic Basin have been analysed to find possible links between variability in the climatic parameters and the phenomenon of mucilage. Seasonal anomalies of temperature, calculated as spatial mean over the Po Valley area, and anomalies of North Atlantic Oscillation were compared with the historical record of mucilage episodes. Both climatic indices were found to be positively correlated with mucilage events, suggesting a possible relationship between climatic variability and the increased appearance of mucilage aggregates.     

A quantitative method for assessing the toxic effects of some water soluble substances, based on changes in periphyton community structure

A method to assess the potential damage of a particular water soluble compound when released in lakes and streams is described. The results of a test using germanium dioxide which has a known effect, provided evidence that quantitative changes in the species composition of the colonizing periphyton community reflect the nature of the toxicant to which they were exposed. Natural diatom growth in the presence of germanium was inhibited and the ensueing changes in the species composition of the treated community were reflected in changes in the index of diversity calculated for that community. The advantages and disadvantages of this approach are discussed.     

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.