We present an overview of the plankton studies conducted during the last 25 years in the epipelagic offshore waters of the Mediterranean Sea. This quasi-enclosed sea is characterized by a rich and ...complex physical dynamics with distinctive traits, especially in regard to the thermohaline circulation. Recent investigations have basically confirmed the long-recognised oligotrophic nature of this sea, which increases along both the west-east and the north-south directions. Nutrient availability is low, especially for phosphorous (N:P up to 60), though this limitation may be buffered by inputs from highly populated coasts and from the atmosphere. Phytoplankton biomass, as chl a, generally displays low values (less than 0.2 μg chl a l−1) over large areas, with a modest late winter increase. A large bloom (up to 3 μg l−1) is observed throughout the late winter and spring exclusively in the NW area. Relatively high biomass values are recorded in fronts and cyclonic gyres. A deep chlorophyll maximum is a permanent feature for the whole basin, except during the late winter mixing. It is found at increasingly greater depths ranging from 30 m in the Alboran Sea to 120 m in the easternmost Levantine basin. Primary production reveals a west-east decreasing trend and ranges between 59 and 150 g C m−2 y−1 (in situ measurements). Overall, the basin is largely dominated by small autotrophs, microheterotrophs and egg-carrying copepod species. The microorganisms (phytoplankton, viruses, bacteria, flagellates and ciliates) and zooplankton components reveal a considerable diversity and variability over spatial and temporal scales, although the latter is poorly studied. Examples are the wide diversity of dinoflagellates and coccolithophores, the multifarious role of diatoms or picoeukaryotes, and the distinct seasonal or spatial patterns of the species-rich copepod genera or families which dominate the basin. Major dissimilarities between western and eastern basins have been highlighted in species composition of phytoplankton and mesozooplankton, but also in the heterotrophic microbial components and in their relationships. Superimposed to these longitudinal differences, a pronounced biological heterogeneity is also observed in areas hosting deep convection, fronts, cyclonic and anti-cyclonic gyres or eddies. In such areas, the intermittent nutrient enrichment promotes a switching between a small-sized microbial community and diatom-dominated populations. A classical food web readily substitutes the microbial food web in these cases. These switches, likely occurring within a continuum of trophic pathways, may greatly increase the flux towards higher trophic levels, in spite of the apparent heterotrophy. Basically, the microbial system seems to be both bottom-up and top-down controlled. A "multivorous web" is shown by the great variety of feeding modes and preferences and by the significant and simultaneous grazing impact on phytoplankton and ciliates by mesozooplankton.
Microbial food web dynamics were determined during the onset of several spring phytoplankton blooms induced by natural iron fertilization off Kerguelen Island in the Southern Ocean (KEOPS2). The ...abundances of heterotrophic bacteria and heterotrophic nanoflagellates, bacterial heterotrophic production, bacterial respiration, and bacterial growth efficiency, were consistently higher in surface waters of the iron-fertilized sites than at the reference site in HNLC (high nutrient low chlorophyll) waters. The abundance of virus-like particles remained unchanged, but viral production increased by a factor of 6 in iron-fertilized waters. Bacterial heterotrophic production was significantly related to heterotrophic nanoflagellate abundance and viral production across all sites, with bacterial production explaining about 70 and 85%, respectively, of the variance of each in the mixed layer (ML). Estimated rates of grazing and viral lysis, however, indicated that heterotrophic nanoflagellates accounted for a substantially higher loss of bacterial production (50%) than viruses (11%). Combining these results with rates of primary production and export determined for the study area, a budget for the flow of carbon through the microbial food web and higher trophic levels during the early (KEOPS2) and the late phase (KEOPS1) of the Kerguelen bloom is provided.
Microbial eukaryotic community composition was examined by 18S rRNA gene tag pyrosequencing, during the early phase of spring phytoplankton blooms induced by natural iron fertilization, off Kerguelen ...Island in the Southern Ocean (KEOPS2 cruise). A total of 999 operational taxonomical units (OTUs), affiliated to 30 known high-level taxonomic groups, were retrieved from 16 samples collected in the upper 300 m water column. The alveolata group was the most abundant in terms of sequence number and diversity (696 OTUs). The majority of alveolata sequences were affiliated to Dinophyceae and to two major groups of marine alveolates (MALV-I and MALV-II). In the upper 180 m, only 13% of the OTUs were shared between of the fertilized stations and the reference site characterized by high-nutrient low-chlorophyll (HNLC) waters. Fungi and Cercozoa were present in iron-fertilized waters, but almost absent in the HNLC samples, while Haptophyta and Chlorophyta characterized the HNLC sample. Finally, the 300 m depth samples of all stations were differentiated by the presence of MALV-II and Radiolaria. Multivariate analysis, examining the level of similarity between different samples, showed that protistan assemblages differed significantly between the HNLC and iron-fertilized stations, but also between the diverse iron-fertilized blooms.
Above the Kerguelen Plateau in the Southern Ocean natural iron fertilization sustains a large phytoplankton bloom over 3 months during austral summer. During the KEOPS1 project (KErguelen Ocean and ...Plateau compared Study1) we sampled this phytoplankton bloom during its declining phase along with the surrounding high-nutrient-low-chlorophyll (HNLC) waters to study the effect of natural iron fertilization on the role of viruses in the microbial food web. Bacterial and viral abundances were 1.7 and 2.1 times, respectively, higher within the bloom than in HNLC waters. Viral production and virus-mediated mortality of bacterioplankton were 4.1 and 4.9 times, respectively, higher in the bloom, while the fraction of infected cells (FIC) and the fraction of lysogenic cells (FLC) showed no significant differences between environments. The present study suggests viruses to be more important for bacterial mortality within the bloom and dominate over grazing of heterotrophic nanoflagellates (HNFs) during the late bloom phase. As a consequence, at least at a late bloom stage, viral lysis shunts part of the photosynthetically fixed carbon in iron-fertilized regions into the dissolved organic matter (DOM) pool with potentially less particulate organic carbon transferred to larger members of the food web or exported.
We investigated the identity of the limiting nutrient of the pelagic microbial food web in the Mediterranean Sea using nutrient manipulated microcosms during summer 2008. Experiments were carried out ...with surface waters at the center of anticyclonic eddies in the Western Basin, the Ionian Basin, and the Levantine Basin. In situ, the ratio of N to P was always higher in both dissolved and particulate organic fractions compared to the Redfield ratio, suggesting a relative P-starvation. In each experiment, four different treatments in triplicates (addition of ammonium, phosphate, a combination of both, and the unamended control) were employed and chemical and biological parameters monitored throughout a 3-4 day incubation. Temporal changes of turnover time of phosphate and ATP, and alkaline phosphatase activity during the incubation suggested that the phytoplankton and heterotrophic prokaryotes (Hprok) communities were not P-limited at the sites. Furthermore, statistical comparison among treatments at the end of the incubation did not support a hypothesis of P-limitation at the three study sites. In contrast, primary production was consistently limited by N, and Hprok growth was not limited by N nor P in the Western Basin, but N-limited in the Ionian Basin, and N and P co-limited in the Levantine Basin. Our results demonstrated the gap between biogeochemical features (an apparent P-starved status) and biological responses (no apparent P-limitation). We question the general notion that Mediterranean surface waters are limited by P alone during the stratified period.
We report on the zonal variability of mesopelagic particulate organic carbon remineralization and deep carbon transfer potential during the Kerguelen Ocean and Plateau compared Study 2 expedition ...(KEOPS 2; October-November 2011) in an area of the polar front supporting recurrent massive blooms from natural Fe fertilization. Mesopelagic carbon remineralization (MR) was assessed using the excess, non-lithogenic particulate barium (Baxs) inventories in mesopelagic waters and compared with bacterial production (BP), surface primary production (PP) and export production (EP). Results for this early season study are compared with the results obtained during a previous study (2005; KEOPS 1) for the same area at a later stage of the phytoplankton bloom. Our results reveal the patchiness of the seasonal advancement and of the establishment of remineralization processes between the plateau (A3) and polar front sites during KEOPS 2. For the Kerguelen plateau (A3 site) we observe a similar functioning of the mesopelagic ecosystem during both seasons (spring and summer), with low and rather stable remineralization fluxes in the mesopelagic column (150-400 m). The shallow water column (~500 m), the lateral advection, the zooplankton grazing pressure and the pulsed nature of the particulate organic carbon (POC) transfer at A3 seem to drive the extent of MR processes on the plateau. For deeper stations (>2000 m) located on the margin, inside a polar front meander, as well as in the vicinity of the polar front, east of Kerguelen, remineralization in the upper 400 m in general represents a larger part of surface carbon export. However, when considering the upper 800 m, in some cases, the entire flux of exported carbon is remineralized. In the polar front meander, where successive stations form a time series, two successive events of particle transfer were evidenced by remineralization rates: a first mesopelagic and deep transfer from a past bloom before the cruise, and a second transfer expanding at mesopelagic layers during the cruise. Regarding the deep carbon transfer efficiency, it appeared that above the plateau (A3 site) the mesopelagic remineralization was not a major barrier to the transfer of organic matter to the seafloor (close to 500 m). There, the efficiency of carbon transfer to the bottom waters (>400 m) as assessed by PP, EP and MR fluxes comparisons reached up to 87% of the carbon exported from the upper 150 m. In contrast, at the deeper locations, mesopelagic remineralization clearly limited the transfer of carbon to depths of >400 m. For sites at the margin of the plateau (station E-4W) and the polar front (station F-L), mesopelagic remineralization even exceeded upper 150 m export, resulting in a zero transfer efficiency to depths >800 m. In the polar front meander (time series), the capacity of the meander to transfer carbon to depth >800 m was highly variable (0 to 73%). The highest carbon transfer efficiencies in the meander are furthermore coupled to intense and complete deep (>800 m) remineralization, resulting again in a near-zero, deep (>2000 m) carbon sequestration efficiency there.
The abundance and activity of the major members of the heterotrophic microbial community – from viruses to ciliates – were studied along a longitudinal transect across the Mediterranean Sea in the ...summer of 2008. The Mediterranean Sea is characterized by a west to-east gradient of deepening of DCM (deep chlorophyll maximum) and increasing oligotrophy reflected in gradients of biomass and production. However, within this well documented longitudinal trend, hydrological mesoscale features exist and likely influence microbial dynamics. Here we present data from a W-E transect of 17 stations during the period of summer stratification. Along the transect the production and fate of organic matter was investigated at three selected sites each one located in the centre of an anticyclonic eddy: in the Algero-Provencal Basin (St. A), the Ionian Basin (St. B), and the Levantine Basin (St. C). The 3 geographically distant eddies showed low values of the different heterotrophic compartments of the microbial food web, and except for viruses in site C, all integrated (0–150 m) stocks were higher in reference stations located in the same basin outside the eddies. During our study the 3 eddies showed equilibrium between GPP (Gross Primary Production) and DCR (Dark Community Respiration). Integrated PPp (Particulate Primary Production) values at A, B and C varied from ~140 to ~190 mg C m−2.
The dilution technique was used to investigate microzooplankton grazing and phytoplankton growth in the eastern English Channel during the diatom–
Phaeocystis spring succession from January 2009 to ...June 2009. Four periods were defined based on phytoplankton composition: Periods 1, 2 and 4 composed of distinct diatom communities of small (5–20
μm length) to larger cells (20–120
μm) in colonies; Period 3 characterized by the
Phaeocystis globosa bloom. Dilution experiments were conducted before, during and after the
P. globosa bloom. Microzooplankton carbon consumption (from 18.1 to 360.9
μg
C L
−
1
d
−
1
) often equalled or exceeded phytoplankton production (from 1.7 to 129.0
μg
C L
−
1
d
−
1
) in particular at the end of the
P. globosa bloom when microzooplankton grazed on previously formed phytoplankton biomass. Results of size-fractionated dilution experiments, conducted with distinct grazer communities, suggested different roles for ciliates and dinoflagellates. Ciliates appeared to be very efficient grazers of small diatoms (5–10
μm) and
P. globosa free cells, whereas dinoflagellates grazed on both larger diatoms (>
10
μm; P1) and small
P. globosa colonies. Ciliates and dinoflagellates did not seem to compete for food resources, as they were oriented towards different phytoplankton size classes.
► Herbivory during the spring phytoplankton bloom in the eastern English Channel. ► 12 dilution experiments carried out with total and size-fractionated Chl
a. ► Experiments were run with different predator/prey communities. ► Ciliates are likely consumers of free cells of
Phaeocystis globosa. ► Dinoflagellates are likely consumers of
Phaeocytis colonies and diatoms.
The community structures and succession of phytoplankton, protozooplankton and copepods were studied from February 2007 to July 2009 in a coastal area of the eastern English Channel subject to ...Phaeocystis globosa blooms. While diatom blooms preceded P. globosa blooms each year, the community structure and stock of heterotrophic protists appeared to be related to the dominant P. globosa life cycle stages. In 2007, the dominance of large colonies (>100 mu m, up to 316 mu g C L super(-1)), which resulted in a high biomass of healthy free cells (up to 132 mu g C L super(-1)), accompanied high spirotrich ciliate stocks (up to 58 mu g C L super(-1)) and high abundances of the copepods Acartia clausi and Temora longicornis (up to 11 ind. L super(-1)). In 2008, the bloom which lasted a shorter period of time was dominated by large colonies (up to 328 mu g C L super(-1)) and fewer free cells (up to 98 mu g C L super(-1)). This corresponded with a lower abundance of grazers, with stocks of heterotrophic protists and copepods 1.6 times and 2.2 times lower, respectively. In 2009, the P. globosa bloom was again dominated by large colonies and <100 mu m diatoms. This corresponded to a dominance of heterotrophic dinoflagellates among the protists (62% of the total heterotrophic protist biomass) and Acartia clausi (55% of the copepod abundance). Overall, heterotrophic dinoflagellates appeared to be likely the most important group of phytoplankton grazers.
The effect of phosphate (P), nitrate (N), and organic carbon (C, glucose) enrichment on heterotrophic bacterial production was examined along two longitudinal transects covering the whole ...Mediterranean Sea during June and September 1999. During these cruises, integrated bacterial production ranged from 11 to 349 mgC m-2d-1for the 0-150 m layer. P was found to stimulate bacterial production (BP) in 13 out of 18 experiments, in the eastern and in the western Mediterranean Sea. Organic carbon stimulation of bacterial production was observed at two stations in the Alboran Sea, where the highest bacterial production was recorded (216 and 349 mg C m-2d-1) and in the Sicily Strait. Maximum rates of alkaline phosphatase (AP) increased from the Alboran to the Levantine Sea whereas AP turnover time decreased. Moreover, alkaline phosphatase activity was not systematically reduced following additions of P. In cases of P limitation, however, the alkaline phosphatase activity to bacterial production ratio was severely reduced in the P and NPC enrichments. Generally, the addition of the limiting factor-whether P or C-had a synchronous stimulating effect on bacterial production and ectoaminopeptidase activity and induced a decline in the amino acid respiration percentage. At two selected stations in the eastern and northwestern Mediterranean, response to enrichment was tested on vertical profiles. Bacteria shifted from P to C limitation at a depth where soluble reactive phosphorus was still undetectable, but corresponding to a strong increase in alkaline phosphatase turnover time. Our results showed that values of AP turnover time lower than 100 h corresponded to situations of P limitation of bacterial production.
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