Emerging concepts on microbial processes in the bathypelagic ocean – ecology,biogeochemistry, and genomics

This paper synthesizes recent findings regarding microbial distributions and processes in the bathypelagic ocean (depth >1000 m). Abundance, production and respiration of prokaryotes reflect supplies of particulate and dissolved organic matter to the bathypelagic zone. Better resolution of carbon fluxes mediated by deep microbes requires further testing on the validity of conversion factors. Archaea, especially marine Crenarchaeota Group I, are abundant in deep waters where they can fix dissolved inorganic carbon. Viruses appear to be important in the microbial loop in deep waters, displaying remarkably high virus to prokaryote abundance ratios in some oceanic regions. Sequencing of 18S rRNA genes revealed a tremendous diversity of small-sized protists in bathypelagic waters. Abundances of heterotrophic nanoflagellates (HNF) and ciliates decrease with depth more steeply than prokaryotes; nonetheless, data indicated that HNF consumed half of prokaryote production in the bathypelagic zone. Aggregates are important habitats for deep-water microbes, which produce more extracellular enzymes (on a per-cell basis) than surface communities. The theory of marine gel formation provides a framework to unravel complex interactions between microbes and organic polymers. Recent data on the effects of hydrostatic pressure on microbial activities indicate that bathypelagic microbial activity is generally higher under in situ pressure conditions than at atmospheric pressures. High-throughput sequencing of 16S rRNA genes revealed a remarkable diversity of Bacteria in the bathypelagic ocean. Metagenomics and comparative genomics of piezophiles reveal not only the high diversity of deep sea microbes but also specific functional attributes of these piezophilic microbes, interpreted as an adaptation to the deep water environment. Taken together, the data compiled on bathypelagic microbes indicate that, despite high-pressure and low-temperature conditions, microbes in the bathypelagic ocean dynamically interact with complex mixtures of organic matter, responding to changes in the ocean’s biogeochemical state.     

Vertical changes in abundance,biomass and community structure of copepods down to 3000 m in the southern Bering Sea

Vertical changes in abundance, biomass and community structure of copepods down to 3000 m depth were studied at a single station of the Aleutian Basin of the Bering Sea (53°28′N, 177°00′W, depth 3779 m) on the 14th June 2006. Both abundance and biomass of copepods were greatest near the surface layer and decreased with increase in depth. Abundance and biomass of copepods integrated over 0–3000 m were 1,390,000 inds. m^?2 and 5056 mg C m^?2, respectively. Copepod carcasses occurred throughout the layer, and the carcass:living specimen ratio was the greatest in the oxygen minimum layer (750–100 m, the ratio was 2.3). A total of 72 calanoid copepod species belonging to 34 genera and 15 families occurred in the 0–3000 m water column (Cyclopoida, Harpacticoida and Poecilostomatoida were not identified to species level). Cluster analysis separated calanoid copepod communities into 5 groups (A–E). Each group was separated by depth, and the depth range of each group was at 0–75 m (A), 75–500 m (B), 500–750 m (C), 750–1500 m (D) and 1500–3000 m (E). Copepods were divided into four types based on the feeding pattern: suspension feeders, suspension feeders in diapause, detritivores and carnivores. In terms of abundance the most dominant group was suspension feeders (mainly Cyclopoida) in the epipelagic zone, and detritivores (mainly Poecilostomatoida) were dominant in the meso- and bathypelagic zones. In terms of biomass, suspension feeders in diapause (calanoid copepods Neocalanus spp. and Eucalanus bungii) were the major component (ca. 10–45%), especially in the 250–3000 m depth. These results are compared with the previous studies in the same region and that down to greater depths in the worldwide oceans.     

Viral abundance and distribution in mesopelagic and bathypelagic waters of the Mediterranean Sea

Despite the fact that marine viruses have been increasingly investigated in the last decade, knowledge on virus abundance, biomass and distribution in mesopelagic and bathypelagic waters is limited. We report here the results of a large-spatial-scale study (covering more than 3000 km) on the virioplankton distribution in epi-, meso- and bathypelagic waters in 19 areas of the Mediterranean Sea, from the Alboran Sea and Western Mediterranean, to the Tyrrhenian Sea, Sicily Channel and Ionian Sea. Integrated viral abundance in epipelagic waters was significantly higher than in deep-sea waters (on average, 2.4 vs. 0.5×10¹² viruses m⁻³). However, abundance of viruses in the deep-Mediterranean waters was the highest reported so far for deep seas worldwide (7.0 and 3.1×10¹¹ viruses m⁻³ in mesopelagic and bathypelagic waters, respectively) and their biomass accounted for 13–18% of total prokaryotic C biomass. The significant relationship between viral abundance and prokaryotic abundance and production in deep waters suggests that also deep-sea viruses are closely dependent on the abundance and metabolism of their hosts. Moreover, virus to prokaryote (and nucleoid-containing cell (NuCC)) abundance ratio increased with increasing depths suggesting that deep waters may represent optimal environments for viral survival or proliferation. Overall, our results indicate that deep waters may represent a significant reservoir of viruses and open new perspectives for future investigations of viral impact on the functioning of meso-bathypelagic ecosystems.     

Diel variations of the bathymetric distribution of zooplankton groups and biomass in Cap-Ferret Canyon,France

The bathymetric distribution, abundance and diel vertical migrations (DVM) of zooplankton were investigated along the axis of the Cap-Ferret Canyon (Bay of Biscay, French Atlantic coast) by a consecutive series of synchronous net hauls that sampled the whole water column (0–2000 m in depth) during a diel cycle. The distribution of appendicularians (maximum 189 individuals m⁻³), cladocerans (maximum 287 individuals m⁻³), copepods (copepods<4 mm, maximum 773 individuals m⁻³, copepods>4 mm, maximum 13 individuals m⁻³), ostracods (maximum 8 individuals m⁻³), siphonophores (maximum >2 individuals m⁻³) and peracarids (maximum >600 individuals 1000 m⁻³) were analysed and represented by isoline diagrams. The biomass of total zooplankton (maximum 18419 μg C m⁻³, 3780 μg N m⁻³) and large copepods (>4 mm maximum 2256 μg C m⁻³, 425 μg N m⁻³) also were determined. Vertical migration was absent or affected only the epipelagic zone for appendicularians, cladocerans, small copepods and siphonophores. Average amplitude of vertical migration was about 400–500 m for ostracods, some hyperiids and mysids, and large copepods, which were often present in the epipelagic, mesopelagic, and bathypelagic zones. Large copepods can constitute more than 80% of the biomass corresponding to total zooplankton. They may play an important role in the active vertical transfer of carbon and nitrogen.     

Microzooplankton composition,biomass and grazing rates along the WOCE SR3 line between Tasmania and Antarctica

Microzooplankton species composition and grazing rates on phytoplankton were investigated along a transect between ∼46 and 67°S, and between 140 and 145°E. Experiments were conducted in summer between November 2nd and December 14th in 2001. The structure of the microbial food web changed considerably along the transect and was associated with marked differences in the physical and chemical environment encountered in the different water masses and frontal regions. On average microzooplankton grazing experiments indicated that 91%, 102%, and 157%, (see results) of the phytoplankton production would be grazed in the <200, <20 and <2 μm size fractions, respectively, indicating microzooplankton grazing was potentially constraining phytoplankton populations (<200 μm) along most of the transect. Small ciliates in general and especially oligotrich species declined in importance from the relatively warm, Southern Subtropical Front waters (6.8 μg C/L) to the colder waters of the southern branch of the Polar Front (S-PF), (∼0.5 μg C/L) before increasing again near the Antarctic landmass. Large changes in microzooplankton dominance were observed, with heterotrophic nanoflagellates (HNF), ciliates and larger dinoflagellates having significant biomass in different water masses. HNF were the dominant grazers when chlorophyll a was low in areas such as the Inter-Polar Frontal Zone (IPFZ), while in areas of elevated biomass such as the S-PF and Southern Antarctic Circumpolar Current (SACC), a mix of copepod nauplii and large heterotrophic and mixotrophic dinoflagellates tended to dominate the grazing community. In the S-PF and SACC water masses the tight coupling observed between the microzooplankton grazers and phytoplankton populations over most of the rest of the transect was relaxed. In these regions grazing was low on the >20 μm size fraction of chlorophyll a, which dominated the biomass, while smaller diatoms and nanoplankton in the <20 μm size fraction were still heavily grazed. The lack of grazing pressure on large phytoplankton contributes to this region's potential to export carbon with larger cells known to have higher sinking rates.     

Abundance,biomass and composition of the sea ice biota of the Greenland Sea pack ice

During two cruises to the Greenland Sea, we studied the abundance and biomass of the sea ice biota in summer and late autumn. The mean calculated biomass of the sympagic community was 0.2 g C m⁻² ice. Algae contributed on average 43% to total biomass, followed by bacteria (31%), heterotrophic flagellates (20%), and meiofauna (4%). Diatoms were the main primary producers (60% of total algal biomass), but flagellated cells contributed significantly to the algal biomass. Among the meiofauna, ciliates, nematodes, acoel turbellarians and crustaceans were dominant. Calculated potential ingestion rates of meiofauna (0.6 g C m⁻² (120 d)⁻¹) are on the same order of magnitude as annual primary production estimates for Arctic multi-year sea ice. We therefore assume that grazing can control biomass accumulation of primary producers inside the sea ice.     

Dissolved organic carbon export and subsequent remineralization in the mesopelagic and bathypelagic realms of the North Atlantic basin

Dissolved organic carbon (DOC) data are presented from three meridional transects conducted in the North Atlantic as part of the US Climate Variability (CLIVAR) Repeat Hydrography program in 2003. The hydrographic sections covered a latitudinal range of 6°S to 63°N along longitudes 20°W (CLIVAR line A16), 52°W (A20) and 66°W (A22). Over 3700 individual measurements reveal unprecedented detail in the DOC distribution and systematic variations in the mesopelagic and bathypelagic zones of the North Atlantic basin. Latitudinal gradients in DOC concentrations combined with published estimates of ventilation rates for the main thermocline and North Atlantic Deep Water (NADW) indicate a net DOC export rate of 0.081 Pg C yr⁻¹ from the epipelagic zone into the mesopelagic and bathypelagic zones. Model II regression and multiple linear regression models applied to pairwise measures of DOC and chlorofluorocarbon (CFC-12) ventilation age, retrieved from major water masses within the main thermocline and NADW, indicate decay rates for exported DOC ranging from 0.13 to 0.94 μmol kg⁻¹ yr⁻¹, with higher DOC concentrations driving higher rates. The contribution of DOC oxidation to oxygen consumption ranged from 5 to 29% while mineralization of sinking biogenic particles drove the balance of the apparent oxygen utilization.     

Major contribution of autotrophy to microbial carbon cycling in the deep North Atlantic’s interior

Current estimates point to a mismatch of particulate organic carbon supply derived from the surface ocean and the microbial organic carbon demand in the meso- and bathypelagic realm. Based on recent findings that chemoautotrophic Crenarchaeota are abundant in the mesopelagic zone, we quantified dissolved inorganic carbon (DIC) fixation in the meso- and bathypelagic North Atlantic and compared it with heterotrophic microbial activity. Measuring ¹⁴C-bicarbonate fixation and ³H-leucine incorporation revealed that microbial DIC fixation is substantial in the mesopelagic water masses, ranging from 0.1 to 56.7 μmol C m⁻³ d⁻¹, and is within the same order of magnitude as heterotrophic microbial activity. Integrated over the dark ocean’s water column, DIC fixation ranged from 1–2.5 mmol C m⁻² d⁻¹, indicating that chemoautotrophy in the dark ocean represents a significant source of autochthonously produced ‘new organic carbon’ in the ocean’s interior amounting to about 15–53% of the phytoplankton export production. Hence, chemoautotrophic DIC fixation in the oxygenated meso- and bathypelagic water column of the North Atlantic might substantially contribute to the organic carbon demand of the deep-water microbial food web.     

Size-fractionated biological iron and carbon uptake along a coastal to offshore transect in the NE Pacific

Iron has been shown to limit phytoplankton growth in high-nutrient low-chlorophyll (HNLC) regions such as the NE subarctic Pacific. We report size-fractionated Fe-uptake rates by the entire plankton community in short (6–8 h) light and dark incubations along an E–W transect from P04 (a coastal ocean station) to OSP (an open-ocean HNLC station) during August–September 1997. Size-fractionated primary productivity and chl a were measured to monitor algal Fe : C uptake ratios and Fe-uptake relative to phytoplankton biomass. The >5.0 μm size-class, which consisted mostly of large diatoms, had the highest Fe-uptake rate at nearshore stations (P04 and P8), but Fe-uptake rates for this size class decreased despite increases in biomass and primary productivity when transecting westwards to HNLC waters. Fe-uptake rates of the small size class (0.2–1.0 μm, including heterotrophic bacteria and autotrophs) were inversely related to the >5.0 μm size-class uptake rates, in that stations with high dissolved Fe (DFe) concentrations had relatively low uptake rates compared to those in the low-Fe offshore region. The 1.0–5.0 μm size-class Fe-uptake rates were low, relatively invariant along the transect, and differed little between light and dark incubations. Dark Fe-uptake rates averaged 10–20% less than those in the light for the >5.0 μm size class. Dark uptake rates however, were higher than light uptake rates for the 0.2–1.0 μm size class at all stations. Fe : C uptake ratios were high for all size classes at P04, but decreased as DFe concentrations decreased offshore. The prokaryote-dominated 0.2–1.0 μm size class had the highest Fe : C uptake ratios at all stations. These data suggest that prokaryotic organisms make an important contribution to biological Fe uptake in this region. Our experiments support the results of previous culture work, suggesting higher Fe : C ratios in coastal phytoplankton compared to open-ocean species, and demonstrate that light can have a large effect on Fe partitioning between size classes in subarctic Pacific HNLC waters.     

Vertical distribution of prokaryote production and abundance in the mesopelagic and bathypelagic layers of the Canada Basin,western Arctic: Implications for the mode and extent of organic carbon delivery

The vertical distributions of prokaryote heterotrophic production (³H-leucine incorporation rate) and abundance were investigated in the meso- and bathy-pelagic layers of the Canada Basin, western Arctic Ocean, during September 2009. Prokaryote production and abundance were high in the Pacific-origin water mass located in the upper mesopelagic layer (depth, 100–200 m). Below the halocline layer (depth, 300–3000 m), both the production and abundance decreased with depth, with log–log regression slopes of −1.33 and −0.77, respectively. Depth-integrated production and biomass in the meso- and bathy-pelagic layers was three- to five-fold lower than the corresponding values reported in the subpolar regions, whereas they were close to or lower than the corresponding values in oligotrophic subtropical regions. Prokaryote turnover times were estimated to be 1.1 and 6.1 years for meso- and bathy-pelagic layers, respectively, with the latter being among the longest turnover times reported for oceanic basins. We estimated prokaryote carbon demand in the water column (100–3000 m) to be on the order of 11 mg C m⁻² d⁻¹, which largely exceeds (by 38-fold) the sinking particulate organic carbon flux at depths of 120–200 m reported in the literature. This large carbon imbalance may be partly explained by organic carbon delivery by lateral intrusion of the Pacific-origin water mass into the upper mesopelagic layer.     

Seasonal changes in zooplankton abundance,biomass, size structure and dominant copepods in the Oyashio region analysed by an optical plankton counter

To identify seasonal patterns of change in zooplankton communities, an optical plankton counter (OPC) and microscopic analysis were utilised to characterise zooplankton samples collected from 0 to 150 m and 0 to 500 m in the Oyashio region every one to three months from 2002 to 2007. Based on the OPC measurements, the abundance and biomass of zooplankton peaked in June (0–150 m) or August (150–500 m), depending on the depth stratum. The peak periods of the copepod species that were dominant in terms of abundance and biomass indicated species-specific patterns. Three Neocalanus species (Neocalanus cristatus, Neocalanus flemingeri and Neocalanus plumchrus) exhibited abundance peaks that occurred before their biomass peaks, whereas Eucalanus bungii and Metridia pacifica experienced biomass peaks before their abundance peaks. The abundance peaks corresponded to the recruitment periods of early copepodid stages, whereas the biomass peaks corresponded to the periods when the dominant populations reached the late copepodid stages (C5 or C6). Because the reproduction of Neocalanus spp. occurred in the deep layer (>500 m), their biomass peaks were observed when the major populations reached stage C5 after the abundance peaks of the early copepodid stages. The reproduction of E. bungii and M. pacifica occurred near the surface layer. These species first formed biomass peaks of C6 and later developed abundance peaks of newly recruited early copepodid stages. From the comparison between OPC measurements and microscopic analyses, seasonal changes in zooplankton biomass at depths of 0–150 m were governed primarily by E. bungii and M. pacifica, whereas those at depths of 150–500 m were primarily caused by the three Neocalanus species.     

Microbial biomass and viral infections of heterotrophic prokaryotes in the sub-surface layer of the central Arctic Ocean

Seawater samples were collected for microbial analyses between 55 and 235 m depth across the Arctic Ocean during the SCICEX 97 expedition (03 September–02 October 1997) using a nuclear submarine as a research platform. Abundances of prokaryotes (range 0.043–0.47×10⁹ dm⁻³) and viruses (range 0.68–11×10⁹ dm⁻³) were correlated (r=0.66, n=150) with an average virus:prokaryote ratio of 26 (range 5–70). Biomass of prokaryotes integrated from 55 to 235 m ranged from 0.27 to 0.85 g C m⁻² exceeding that of phytoplankton (0.005–0.2 g C m⁻²) or viruses (0.02–0.05 g C m⁻²) over the same depth range by an order of magnitude on average. Using transmission electron microscopy (TEM), we estimated that 0.5% of the prokaryote community on average (range 0–1.4%) was visibly infected with viruses, which suggests that very little of prokaryotic secondary production was lost due to viral lysis. Intracellular viruses ranged from 5 to >200/cell, with an average apparent burst size of 45±38 (mean±s.d.; n=45). TEM also revealed the presence of putative metal-precipitating bacteria in 8 of 13 samples, which averaged 0.3% of the total prokaryote community (range 0–1%). If these prokaryotes are accessible to protistan grazers, the Fe and Mn associated with their capsules might be an important source of trace metals to the planktonic food web. After combining our abundance and mortality data with data from the literature, we conclude that the biomass of prokaryoplankton exceeds that of phytoplankton when averaged over the upper 250 m of the central Arctic Ocean and that the fate of this biomass is poorly understood.     

Phytodetritus and the abundance and biomass of abyssal nematodes in the central,equatorial Pacific

The abundance and biomass of abyssal (4300–5000 m) nematodes were investigated along a latitudinal gradient of phytodetritus deposition from 0 to 23°N in the central, equatorial Pacific (140–158°W). Nematode abundance in the oligotrophic, central, North Pacific gyre was 50% lower than at the equator while macrofauna abundance increased 6.5 times over the same transect. Nematode abundance and biomass in the surface (0–1 cm) sediment layer were significantly higher at phytodetritus stations than at non-phytodetritus stations. Abundance and biomass were within the range recorded from other sites of comparable depth that also receive an input of phytodetritus. Abundance was also strongly correlated with microbial biomass. An increase in body size was associated with an increase in food supply. The results demonstrate that the equatorial Pacific represents an immense zone of relatively high nematode standing stock.     

Chemical composition and energy content of deep-sea calanoid copepods in the Western North Pacific Ocean

Condition factor index [CFI=1000×DW/(PL)³; DW: dry weight, PL: prosome length], water content, carbon (C), nitrogen (N), ash and energy content were determined on a total of 69 copepod species caught from the mesopelagic (500–1000 m), upper-bathypelagic (1000–2000 m), lower-bathypelagic (2000–3000 m) and abyssopelagic (3000–5000 m) zones of the western subarctic Pacific. Resultant data were grouped into these four sampling zones, four developmental stage/sex categories (C4, C5 and C6 females and males), three feeding types (carnivore, detritivore and suspension feeder), or two reaction speed groups by the presence/absence of myelinated sheath enveloping axons (fast and slow reacting species). Zone-structured data showed the overall ranges were 3.8–4.6 mm for PL, 1.6–2.6 mg for DW, 21.4–25.0 for CFI, 75.0–78.6% of wet weight (WW) for water, 51.3–53.7% of DW for C, 7.7–8.8% of DW for N, 6.2–7.0 (by weight) for C/N, 6.9–9.6% of DW for ash and 25.3–27.4 J mg⁻¹ DW for energy. Among these components, N and ash exhibited significant between-zone differences characterized by gradual decrease downward for the former, and only the upper-bathypelagic zone>abyssopelagic zone for the latter. Stage/sex-structured data showed no significant differences among them, but energy content of C5 was higher than that of C6 females. From the analyses of feeding type-structured data, carnivores were shown to have lower water, N, ash, but higher C, C/N and energy contents than suspension feeders do. Reaction speed-structured data indicated that slow-reacting species have significantly higher water but lower CFI, C, N and energy contents than fast-reacting species. Designating these grouping criteria, PL and DW as independent variables, the attributes of these variables to the CFI, chemical composition or energy contents were evaluated by stepwise-multiple regression analysis, showing the most pronounced effect of suspension-feeder, followed by the presence of myelinated sheath, DW, C6 females and the abyssopelagic zone. Further analysis of zone-structured data, by adding epipelagic copepod data from identical thermal habitats (Arctic/Antarctic waters), revealed a more marked decline in N content from the epipelagic zone to the abyssopelagic zone, accompanied by an increase in C/N ratios downward. The decline in N (=protein or muscle) contents with depth cannot be explained by the “visual interactions” hypotheses being proposed for the metabolism of pelagic visual predators, but is consistent with the “predation-mediated selection” hypothesis for the metabolism of pelagic copepods.     

Dissolved organic carbon and apparent oxygen utilization in the Atlantic Ocean

Dissolved organic carbon (DOC) distributions along two Atlantic Meridional Transects conducted in 2005 in the region between 47°N and 34°S showed clear latitudinal patterns. The DOC concentrations in the epipelagic zone (0–100 m) were the highest (70–90 µM) in tropical and subtropical waters with stable mixed layers, and lowest (50–55 µM) at the poleward extremities of the transects due to deep convective mixing supplying low DOC waters to the surface. A decrease in DOC occurred with depth, and lowest DOC concentrations (41–45 µM) in the 100–300 m depth range were observed in the equatorial region due to upwelling of low DOC waters. A strong relationship between DOC and AOU was observed in the σ–t 26–26.5 isopycnal layer which underlies the euphotic zone and outcrops at the poleward extremities of the North and South Atlantic Subtropical Gyres (NASG and SASG) in the region ventilating the thermocline waters. Our observations reveal significant north–south variability in the DOC–AOU relationship. The gradient of the relationship suggests that 52% of the AOU in the σ–t 26–26.5 density range was driven by DOC degradation in the NASG and 36% in the SASG, with the remainder due to the remineralisation of sinking particulate material. We assess possible causes for the greater contribution of DOC remineralisation in the NASG compared to the SASG.     

Biogeochemical cycling of N and Si during the mesoscale iron-enrichment experiment in the western subarctic Pacific (SEEDS-II)

Biogeochemical cycles of N and Si were examined in the surface mixed layer during the mesoscale iron-enrichment (IE) experiment in the high-nutrient low-chlorophyll (HNLC) western subarctic Pacific (SEEDS-II). Although the IEs increased nitrate uptake, silicic acid utilization was not stimulated. The nitrate drawdown in the iron-patch (IN-patch, 140.3 mmol m⁻² in the surface mixed layer, 0–30 m) was only 25% of the initial inventory, which was 1/3–2/5 of the previous IE experiments in the subarctic Pacific. This relatively weak response of nutrient drawdown to IEs was due to the high biomass of mesozooplankton (MZ) dominated by copepod Neocalanus plumchrus. Feeding of MZ (247.2 mmol m⁻² during Day 0–21 from the first IE) in the IN-patch was higher than the nitrate drawdown and prevented further development of the phytoplankton bloom. In the later period of the experiment (Day 14–21), the increase in the feeding activity and resultant decrease in phytoplankton biomass induced the accumulation of dissolved organic nitrogen (DON) and ammonium. Among total growth of MZ (81.6 mmol N m⁻²), 89% (72.8 mmol N m⁻²) was transported to the depth by the ontogenetic downward migration of N. plumchrus. Although silicic acid drawdown was not increased by the IEs, Si export flux increased by 2.7 times. The increase in Si export was also due to the increase in MZ, which egested faecal pellets with higher Si:N ratio and faster sinking speed than diatoms. The export efficiency (78% of new production) and total amount of export flux (143.8 mmol N m⁻², 1392 mmol C m⁻²) were highest records within the IE experiments despite weak responses of nutrient drawdown to the IE. During SEEDS-II, the high biomass of MZ reduced the phytoplankton response and nutrient drawdown to the IEs but via grazing and ontogenetic vertical migration accelerated the export flux as well as accumulations of dissolved forms of N. Results of the present and previous IE experiments indicate that the ecosystem and biogeochemical responses to IEs in the HNLC region are quite sensitive to the ecosystem components, especially for grazers of diatoms such as copepods and heterotrophic dinoflagellates. More attention needs to be paid to the ecosystem components and their biogeochemical functions as well as physical and chemical properties of the ecosystems in order to hindcast or forecast the impacts of changes in atmospheric iron deposition.     

Diel vertical distribution of planktonic ciliates within the surface layer of the NW Mediterranean (May 1995)

The composition and vertical distribution of planktonic ciliates within the surface layer was monitored over four diel cycles in May 95, during the JGOFS-France DYNAPROC cruise in the Ligurian Sea (NW Mediterranean). Ciliates were placed into size and trophic categories: micro- and nano-heterotrophic ciliates, mixotrophic ciliates, tintinnids and the autotrophic Mesodinium rubrum. Mixotrophic ciliates (micro and nano) represented an average of 46% of oligotrich abundance and 39% of oligotrich biomass; nano-ciliates (hetero and mixotrophic) were abundant, representing about 60 and 17% of oligotrich abundance and biomass, respectively. Tintinnid ciliates were a minor part of heterotrophic ciliates. The estimated contribution of mixotrophs to chlorophyll a concentration was modest, never exceeding 9% in discrete samples. Vertical profiles of ciliates showed that chlorophyll-containing ciliates (mixotrophs and autotrophs) were mainly concentrated and remained at the chlorophyll a maximum depth. In contrast, among heterotrophic ciliates, a portion of the population appeared to migrate from 20–30 m depth during the day to the surface at night or in the early morning. Correlation analyses of ciliate groups and phytoplankton pigments showed a strong relationship between nano-ciliates and zeaxanthin, and between chlorophyll-containing ciliates and chlorophyll a, as well as other pigments that were maximal at the chlorophyll a maximum depth. Total surface layer concentrations showed minima of ciliates during nightime/early morning hours.     

The structure of zooplankton communities,in the 2 to 2000 μm size range,in the Arabian Sea during and after the SW monsoon, 1994

Zooplankton communities, studied in the surface mixed layer on a 1600 m transect across the Arabian Sea, were found to differ in their temporal and spatial response to seasonal forcing. The transect studied, spanned seasonally eutrophic upwelling, mesotrophic downwelling and aseasonal oligotrophic waters. The nano- and microzooplankton communities constituted a relatively constant compartment in the tropical monsoon ecosystem, whilst the mesozooplankton showed a clear response to both upwelling and season. The heterotrophic nanoflagellates were concentrated in the surface mixed layer, except in the eutrophic upwelling waters of the SW monsoon. They reached maximum cell concentrations of 855 ml^-1 during the SW monsoon and a maximum biomass of 8.4 mg C m^-3 during the intermonsoon. Nanozooplankton standing stocks, in the surface mixed layer, ranged between 7 and 333 mg C m^-2, with highest stocks found during the intermonsoon. The microzooplankton community was dominated by Protozoa, particularly aloricate ciliates and heterotrophic dinoflagellates, which accounted for up to 99% in terms of numbers and up to 71% of the biomass. Sarcodines and metazoan nauplii were recorded in lower numbers (<400 l^-1). The microzooplankton were also concentrated in the surface mixed layer during both periods, except in the eutrophic coastal waters during the SW monsoon, when relatively high biomass values were found below the mixed layer depth. Their standing stocks, in the surface mixed layer, ranged between 50 and 182 mg C m^-2, with the highest concentration found in the mesotrophic offshore waters during the late monsoon period. Total mesozooplankton standing stocks, in the surface 100 m, decreased with distance from the coastal to offshore waters and between seasons, decreasing from 1248 to 238 mg C m^-2 during the late SW monsoon and 656–89 mg C m^-2 during the following intermonsoon. The largest size class, of 1000–2000 μm sized organisms, dominated throughout except at the oligotrophic station during the intermonsoon period, when the smallest class, of 200–500 μm, were more important. The shift in size structure from large to small zooplankton occurred in response to a shift in dominance from large to small phytoplankton cells both spatially, along a eutrophic–oligotrophic gradient, and seasonally. These responses are a result of the physical forcing associated with the monsoon seasons in the Arabian Sea.     

Distribution and relative importance of jellyfish in a region of hydrothermal venting

Net sampling to 3000 m depth at Endeavour Ridge in the northeast Pacific in July 1991–1994 shows that medusae in the immediate vicinity of the hydrothermal vent fields often make up a larger proportion of the total zooplankton abundance and biomass from mesopelagic to bathypelagic depths than in the surrounding waters. This was particularly evident in the dominant Trachymedusae, and least evident in the siphonophores. In addition, the large red Scyphomedusa Stygiomedusa gigantea was a major biomass component in the region of the deep (1000–1800 m depth) migrating scattering layers at the vent field, but was not found in any net tows greater than 10 km away from vents. There is no concurrent increase in relative or percent biomass of fish or chaetognaths, which are the other major predators in the community. We hypothesize that predaceous medusae respond opportunistically to the enhanced zooplankton biomass throughout the water column around vents in spring to early summer, in a way that other predators do not.