On processes determining the vertical stability of surface waters in the marginal ice zone of the north-western Weddell Sea and their relationship with phytoplankton bloom development

Summary Ice-edge-related phytoplankton blooms following the retreating sea-ice in the marginal ice zone are frequently observed phenomena. Such blooms are generally short-lived and are followed by a strong decrease in the chlorophyll concentration towards the open ocean, generally explained by the degradation of the vertical stability. Solar heating and ice melting, which control the stability of the surface water of the north-western Weddell Sea during sea-ice retreat in spring were analysed in order to relate the spatial features of the phytoplankton ice-edge bloom in 1988 to the density field. Solar heating has little effect on the density of seawater in ice-covered areas because the thermal expansion coefficient is very low close to the freezing temperature. Outside the marginal ice zone, the temperature effect on stratification increases and the combined contribution of melting and heating on buoyancy input to the surface layer is roughly constant. As a consequence, the low phytoplankton stocks in the open ocean adjacent to the marginal ice zone, after an initial bloom peak following the retreating ice in spring, are not necessarily caused by deep turbulent mixing, in contrast to what is often assumed.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Temporal and spatial variation of the phytoplankton assemblage in the eastern Indian sector of the Southern Ocean in summer 2001/2002

The abundance and species composition of phytoplankton were investigated at stations in a permanently ice-free (61°S) and seasonally ice-covered area (64°S and 66°30′S) in the eastern Indian sector of the Southern Ocean between November 2001 and March 2002. Although a phytoplankton bloom occurred just after retreat of the sea ice at both stations in the seasonally ice-covered area, vertical stability of the water column during the bloom was weak at the most southerly station. This shows that a bloom can form even under weak vertical stability. In the bloom, diatoms dominated under weak vertical stability and Phaeocystis under strong vertical stability. In the latter case, ice algae largely contributed to development of the bloom. In the later observation period, a subsurface chlorophyll maximum (SCM) was observed at 61°S and 64°S. Species composition was different between the mixed layer and SCM at 64°S, but was uniform with depth at 61°S, indicating that the SCM is formed by different mechanisms.     

Seasonal shifts in ice edge phytoplankton blooms in the Barents Sea related to the water column stability

The development of the phytoplankton bloom and its relation to water column stabilisation during the transition from early to high summer (of 1991) in the seasonally ice-covered zone of the Barents Sea were studied from a meridional transect of repeated hydrographic/biological stations. The water column stabilisation is described in detail with the aid of vertical profiles of the Brunt-Väisälä frequency squared (N²). The contributions of seasonal warming and ice melting to stabilisation are elucidated by determining the effects of temperature and salinity on N². The spring bloom in 1991 migrated poleward from June to July by about 400 km, associated with the retreat of the ice edge. The spring bloom culminated with maximum chlorophyll concentrations in the mixed layer about 100–300 km north of the centre of the meltwater lens, at its northern edge, where the ice cover was still substantial. From the distribution of N² it becomes obvious that the bloom starts at the very beginning of stabilisation, which results solely from the release of meltwater. The increase in temperature due to the seasonal warming does not contribute to the onset of vernal blooming; temperature starts to contribute to the stratification later, when the spring bloom has ceased due to the exhaustion of nutrients in the mixed layer. By that time a deep chlorophyll maximum has formed in the seasonal pycnocline, 20–30 m below the base of the mixed layer. The effect of the seasonal ice cover on the mean areal new primary production is discussed.     

Dynamics of ice algae and phytoplankton in Frobisher Bay

Summary Vertical and seasonal variations of ice algae and phytoplankton were studied in relation to their physico-chemical environments in Frobisher Bay from 1979 to 1986. The biomass, estimated by both chlorophyll a concentrations and cell counts, was greater in the ice algae than in the phytoplankton in the underlying sea-water during winter and spring. Algal distribution in the sea ice varied vertically and seasonally, while in the underlying water column the phytoplankton distribution was much less variable. The ice algal bloom occurred at the bottom of the ice, particularly in the lower 5 cm during late spring, while the phytoplankton bloom took place at depths between 1 and 10 m during early summer after the ice bloom was over. The community structure of the ice algae changed from pennate to centric diatoms as the ice melted. The centrics dominated through the fall, and then decreased as the pennates increased in dominance when the ice formed again in winter. Species diversity and number were greater in the sea ice than in the seawater, but they were similar vertically within each habitat. The evenness of the species distribution did not vary with ice thickness or water depth. Species composition, abundance and dominance of ice algae and phytoplankton continually change both vertically and seasonally. The differential abilities of the species to attain maximal growth rates under various environmental conditions may result in species succession. Evidence is given for the major role of environmental factors regulating the dynamics of ice algae and phytoplankton.     

Abundance,population structure and vertical distribution of dominant calanoid copepods on the eastern Weddell Sea shelf during a spring phytoplankton bloom

The main emphasis of this study was to analyse the short-term development of abundance, population structure and vertical distribution of the dominant calanoid copepods during a phytoplankton bloom in the coastal area of the eastern Weddell Sea in December 2003. Microcalanus pygmaeus was by far the most abundant calanoid species. Metridia gerlachei, Ctenocalanus citer, Calanoides acutus, Calanus propinquus and the ice-associated Stephos longipes were also present in considerable proportions. The observed changes in the population characteristics and parameters of these species are described in detail and discussed in the context of the spring phytoplankton bloom. A conspicuous event occurring during the final stage of the study was the development of a strong storm. While the results suggest that this storm did not have any considerable influence on the populations of all other investigated copepod species, it very likely caused pronounced changes in the S. longipes population present in the water column. Before the storm, S. longipes was found primarily in the upper 100 m of the water column, and its population was dominated by adults (mean proportion = 41%) and the copepodite stage I (mean proportion = 30%). After the storm, the abundance increased considerably, and the copepodite stage I contributed by far the largest proportion (53%) of the total population indicating that the early copepodite stages probably had been released from the sea ice into the under ice water layer due to ice break-up and ice melt processes caused by the storm.     

Bacterial activity in sea ice and open water of the Weddell Sea,Antarctica: A microautoradiographic study

Metabolic activity of bacteria was investigated in open water, newly forming sea ice, and successive stages of pack ice in the Weddell Sea. Microautoradiography, using [³H]leucine as substrate, was compared with incorporation rates of [³H]leucine into proteins. Relation of [³H]leucine incorporation to the biomass of active bacteria provides information about changes of specific metabolic activity of cells. During a phytoplankton bloom in an ice-free, stratified water column, total numbers of bacteria in the euphotic zone averaged 2.3 × 10⁵ ml^–1, but only about 13% showed activity via leucine uptake. Growth rate of the active bacteria was estimated as 0.3–0.4 days^–1. Total cell concentration of bacteria in 400 m depth was 6.6 × 10⁴ ml^–1. Nearly 50% of these cells were active, although biomass production and specific growth rate were only about one-tenth that of the surface populations. When sea ice was forming in high concentrations of phytoplankton, bacterial biomass in the newly formed ice was 49.1 ng C ml^–1, exceeding that in open water by about one order of magnitude. Attachment of large bacteria to algal cells seems to cause their enrichment in the new ice, since specific bacterial activity was reduced during ice formation, and enrichment of bacteria was not observed when ice formed at low algal concentration. During growth of pack ice, biomass of bacteria increased within the brine channel system. Specific activity was still reduced at these later stages of ice development, and percentages of active cells were as low as 3–5%. In old, thick pack ice, bacterial activity was high and about 30% of cells were active. However, biomass-specific activity of bacteria remained significantly lower than that in open water. It is concluded that bacterial assemblages different to those of open water developed within the ice and were dominated by bacteria with lower average metabolic activity than those of ice-free water.     

Factors controlling phytoplankton ice-edge blooms in the marginal ice-zone of the northwestern Weddell Sea during sea ice retreat 1988: Field observations and mathematical modelling

The factors controlling phytoplankton bloom development in the marginal ice zone of the northwestern Weddell Sea were investigated during the EPOS (Leg 2) expedition (1988). Measurements were made of physical and chemical processes and biological activities associated with the process of ice-melting and their controlling variables particularly light limitation mediated by vertical stability and ice-cover, trace metal deficiency and grazing pressure. The combined observations and process studies show that the initiation of the phytoplankton bloom, dominated by nanoplanktonic species, was determined by the physical processes operating in the marginal ice zone at the time of ice melting. The additional effects of grazing pressure by protozoa and deep mixing appeared responsible for a rather moderate phytoplankton biomass (4 mg Chla m^–3) with a relatively narrow geographical extent (100–150 km). The rôle of trace constituents, in particular iron, was minor. The importance of each factor during the seasonal development of the ice-edge phytoplankton bloom was studied through modelling of reasonable scenarios of meteorological and biological forcing, making use of a one-dimensional coupled physicalbiological model. The analysis of simulations clearly shows that wind mixing events — their duration, strength and frequency — determines both the distance from the iceedge of the sea ice associated phytoplankton bloom and the occurrence in the ice-free area of secondary phytoplankton blooms during the summer period. The magnitude and extent of the ice-edge bloom is determined by the combined action of meteorological conditions and grazing pressure. In the absence of grazers, a maximum ice-edge bloom of 7.5 mg Chla m^–3 is predicted under averaged wind conditions of 8 m s^–1. Extreme constant wind scenarios (4–14 m s^–1) combined with realistic grazing pressure predict maximum ice-edge phytoplankton concentrations varying from 11.5 to 2 mg Chla m^–3. Persistent violent wind conditions ( 14 m s^–1) are shown to prevent blooms from developing even during the brightest period of the year.     

A small population of planktonic Flavobacteria with disproportionally high growth during the spring phytoplankton bloom in a prealpine lake

Bacterioplankton growth in temperate Lake Zurich (Switzerland) was studied during the spring phytoplankton bloom by in situ techniques and short-term dilution bioassays. A peak of chlorophyll a (Chl a ) concentrations was followed by a rise of bacterial cell numbers and leucine assimilation rates, of the proportions of cells incorporating 5-bromo-2-deoxyuridine (BrdU), and of community net growth rates in dilution cultures. Incorporation of BrdU was low in Betaproteobacteria (2 ± 1%), indicating that these bacteria did not incorporate the tracer. Pronounced growth of Betaproteobacteria in the enrichments was only observed after the decline of the phytoplankton bloom. An initial peak in the proportions of BrdU-positive Actinobacteria (30%) preceded a distinct rise of their cell numbers during the period of the Chl a maximum. Cytophaga–Flavobacteria (CF) changed little in numbers, but featured high proportions of BrdU-positive cells (28 ± 12%). Moreover, CF represented > 90% of all newly formed cells in dilution cultures before and during the phytoplankton bloom. One phylogenetic lineage of cultivable Flavobacteria (FLAV2) represented a small (0.5–1%) but highly active population in lake plankton. The growth rates of FLAV2 in dilution cultures doubled during the period of the Chl a maximum, indicating stimulation by phytoplankton exudates. Thus, CF, and specifically Flavobacteria , appeared to be substantially more important for carbon transfer in Lake Zurich spring bacterioplankton than was suggested by their standing stocks. The high in situ growth potential of these bacteria might have been counterbalanced by top-down control.     

The impact of sea ice on the initiation of the spring bloom on the Newfoundland and Labrador Shelves

The relationship between sea ice and the phytoplankton springbloom over the Newfoundland and Labrador shelves is examinedusing remotely-sensed chlorophyll data and sea-ice data forthe period 1998–2004. A regression analysis between thetwo data sets shows that the retreat of sea ice precedes thespring bloom, and the inter-annual variation of the spring bloomis closely correlated with the start time of ice retreat. Thespring bloom off Canada's east coast usually starts on the easternGrand Banks. Here, the water properties are strongly influencedby sea ice on the Newfoundland shelves in early spring whenaccelerated ice melting causes the ice edge to retreat northand the melt water is advected south by the Labrador Current.After the ice retreat, the water on the eastern Grand Banksis rapidly stratified and the mixed layer shallows as a resultof surface freshening. The shallow mixed layer promotes phytoplanktongrowth. The regression analysis also reveals that an early springbloom or ice retreat tends to prolong the duration of the springbloom.     

Sedimentation in Arctic Canada: Species composition and biomass of phytoplankton contributed to the marine sediments in Frobisher Bay

Summary Sedimentation of phytoplankton provides food and energy for zoobenthic communities. In this study the rates, species composition and biomass of phytoplankton input to Frobisher Bay sediments were examined during ice (late November to July) and open water (late July to October) periods from 1982 to 1985. The rates were higher on the sea bed than at 20 m. The minimum rate (3x10⁵ cells·m^-2·day^-1) of sedimentation occurred during the early part of the ice period. It increased as the ice thickened and reached a maximum of 2.8x10⁸ cells·m^-2·day^-1 after the phytoplankton bloom at the beginning of the open water period in the first two weeks of August. The sedimented phytoplankton was dominated by diatoms, with a great majority of pennate species during the spring (April to June) and centric forms during the summer (July to August). Green flagellates, dinoflagellates and chrysophytes occurred as a low percentage of the total population in all seasons. Other indicators (chlorophyll a and phaeopigments) showed highest biomass levels in the deepest traps. They were consistently low during the winter (December to March) and reached their maxima during the open-water period of summer. Their abundance was correlated with the seasonal cycle of the phytoplankton in the water column.     

Timing of blooms,algal food quality and Calanus glacialis reproduction and growth in a changing Arctic

The Arctic bloom consists of two distinct categories of primary producers, ice algae growing within and on the underside of the sea ice, and phytoplankton growing in open waters. Long chain omega‐3 fatty acids, a subgroup of polyunsaturated fatty acids (PUFAs) produced exclusively by these algae, are essential to all marine organisms for successful reproduction, growth, and development. During an extensive field study in the Arctic shelf seas, we followed the seasonal biomass development of ice algae and phytoplankton and their food quality in terms of their relative PUFA content. The first PUFA‐peak occurred in late April during solid ice cover at the onset of the ice algal bloom, and the second PUFA‐peak occurred in early July just after the ice break‐up at the onset of the phytoplankton bloom. The reproduction and growth of the key Arctic grazer Calanus glacialis perfectly coincided with these two bloom events. Females of C. glacialis utilized the high‐quality ice algal bloom to fuel early maturation and reproduction, whereas the resulting offspring had access to ample high‐quality food during the phytoplankton bloom 2 months later. Reduction in sea ice thickness and coverage area will alter the current primary production regime due to earlier ice break‐up and onset of the phytoplankton bloom. A potential mismatch between the two primary production peaks of high‐quality food and the reproductive cycle of key Arctic grazers may have negative consequences for the entire lipid‐driven Arctic marine ecosystem.     

Diatom assemblages promote ice formation in large lakes

We present evidence for the directed formation of ice by planktonic communities dominated by filamentous diatoms sampled from the ice-covered Laurentian Great Lakes. We hypothesize that ice formation promotes attachment of these non-motile phytoplankton to overlying ice, thereby maintaining a favorable position for the diatoms in the photic zone. However, it is unclear whether the diatoms themselves are responsible for ice nucleation. Scanning electron microscopy revealed associations of bacterial epiphytes with the dominant diatoms of the phytoplankton assemblage, and bacteria isolated from the phytoplankton showed elevated temperatures of crystallization (T_c) as high as −3 °C. Ice nucleation-active bacteria were identified as belonging to the genus Pseudomonas, but we could not demonstrate that they were sufficiently abundant to incite the observed freezing. Regardless of the source of ice nucleation activity, the resulting production of frazil ice may provide a means for the diatoms to be recruited to the overlying lake ice, thereby increasing their fitness. Bacterial epiphytes are likewise expected to benefit from their association with the diatoms as recipients of organic carbon excreted by their hosts. This novel mechanism illuminates a previously undescribed stage of the life cycle of the meroplanktonic diatoms that bloom in Lake Erie and other Great Lakes during winter and offers a model relevant to aquatic ecosystems having seasonal ice cover around the world.     

Experimental formation of sea ice: importance of water circulation and wave action for incorporation of phytoplankton and bacteria

The efficiency of physical concentration mechanisms for enrichment of algae and bacteria in newly formed sea-ice was investigated under defined conditions in the laboratory. Sea-ice formation was simulated in a 3,000 l tank under different patterns of water movement. When ice formed in an artificially generated current pattern, algal cells were substantially enriched within the ice matrix. Enrichment factors for chlorophyll a calculated from the ratio between the concentrations in ice and underlying water reached values of up to 53. Repeated mixing of ice crystals into the water column, as well as flow of water through the new ice layer, contributed to the enrichment of algae in the ice. Wave action during ice formation revealed lower phytoplankton enrichment factors of up to 9. Mixing of floating ice crystals with underlying water and pumping of water into the ice matrix by periodical expansion and compression of the slush ice layer were responsible for the wave-induced enrichment of algal cells. Physical enrichment of bacteria within the ice was negligible. Bacterial biomass within new ice was enhanced only when the concentration of algae was high. At low algal biomass, bacteria experienced substantial losses in the ice, most likely due to brine drainage, which were not observed for the microalgae. Bacterial cells are therefore not scavenged by ice crystals and the observed enrichment and sustainment of bacterial biomass within newly formed ice depend on their attachment to cells or aggregates of algae. Division rates of bacteria changed only slightly during ice formation. Received: 21 October 1997 / Accepted: 9 April 1998     

Aggregation of algae released from melting sea ice: implications for seeding and sedimentation

Summary Factors influencing the fate of ice algae released from melting sea ice were studied during a R V Polarstern cruise (EPOS Leg 2) to the northwestern Weddell Sea. The large-scale phytoplankton distribution patterns across the receding ice edge and small-scale profiling of the water column adjacent to melting ice floes indicated marked patchiness on both scales. The contribution of typical ice algae to the phytoplankton was not significant. In experiments simulating the conditions during sea ice melting, ice algae revealed a strong propensity to form aggregates. Differences in the aggregation potential were found for algal assemblages collected from the ice interior and the infiltration layer. Although all algal species collected from the ice were also found in aggregates, the species composition of dispersed and aggregated algae differed significantly. Aggregates were of a characteristic structure consisting of monospecific microaggregates which are likely to have formed in the minute brine pockets and channels within the ice. Sinking rates of aggregates were three orders of magnitude higher than those of dispersed ice algae. These observations, combined with the negligible seeding effect of ice algae found during this study, suggest that ice algae released from the melting sea ice are subject to rapid sedimentation. High grazing pressure at the ice edge of the investigation area is another factor eliminating ice algae released during melting.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Annual variability in ciliate community structure, potential prey and predators in the open northern Baltic Sea proper

Biomass of ciliates, bacteria and mesozooplankton, as well asbiomass estimates of phytoplankton from chlorophyll a values,were studied in the mixed layer of the northern Baltic Sea proper,between February and December 1998. Production of phytoplanktonand bacteria was measured, and production of ciliates and mesozooplanktonwas estimated. The phytoplankton spring bloom in late Marchwas dominated by diatoms and dinoflagellates. Ciliates had abiomass peak shortly after the spring bloom, while mesozooplanktonpeaked in July. Thus, the predation pressure on ciliates waslow in spring, and ciliates were major predators, potentiallyconsuming up to 15% of the primary production. In summer, therewas a shift from larger to smaller ciliates coinciding witha shift from larger to smaller primary producers, an increasein bacterial production, and also an increase in mesozooplanktonabundance, mainly copepods. Elevated mesozooplankton predationand selective removal of larger ciliate species and/or a shiftto smaller prey size presumably caused these changes. The potentialcarbon consumption from ciliates and mesozooplankton was highestin summer and autumn, reaching 55 and 40% of the primary productionin summer and autumn, respectively. Ciliates consumed twiceas much as mesozooplankton, thus acting as important regenerators.     

Phytoplankton in the Marginal Ice Zone of the Greenland Sea during summer, 1984

Summary Phytoplankton biomass and species composition were studied in transects through the ice edge region of the Greeland Sea from 19 July to 8 August 1984. Biomass was estimated by vertical in situ chlorophyll fluorescence and pigment extraction of discrete samples. Preserved material was used for identification of phytoplankton species and calculation of their relative abundances. The results suggest that the various geographical regions of the Greenland Sea differ considerably in their phytoplankton development. Autotrophic biomass and species composition were closely associated with the extent of the annual and seasonal ice cover, hydrographic conditions, nutrient availability and the water masses typical of the different domains. In the NE Greenland polynya a deep mixed layer inhibited the development of a phytoplankton bloom, whereas greatest biomass concentrations were associated with a receding ice edge on the E Greenland Shelf. In the Fram Strait, the position of the relatively stationary ice edge is controlled by frontal dynamics, currents and wind. Due to rapidly changing physical and chemical conditions, phytoplankton biomass showed great variability between stations. High chlorophyll a concentrations may develop locally where melting ice causes stratification or can result from passive accumulation in eddies. In July/August 84 the Fram Strait area was dominated by a typical summer population of flagellates and large diatom species.Contribution 6 of the Alfred-Wegener-Institute for Polar and Marine Research     

Cultivation and ecosystem role of a marine roseobacter clade-affiliated cluster bacterium

Isolation and cultivation are a crucial step in elucidating the physiology, biogeochemistry, and ecosystem role of microorganisms. Many abundant marine bacteria, including the widespread Roseobacter clade-affiliated (RCA) cluster group, have not been cultured with traditional methods. Using novel techniques of cocultivation with algal cultures, we have accomplished successful isolation and propagation of a strain of the RCA cluster. Our experiments revealed that, in addition to growing on alga-excreted organic matter, additions of washed bacterial cells led to significant biomass decrease of dinoflagellate cultures as measured by in vivo fluorescence. Bacterial filtrate did not adversely affect the algal cultures, suggesting attachment-mediated activity. Using an RCA cluster-specific rRNA probe, we documented increasing attachment of these algicidal bacteria during a dinoflagellate bloom, with a maximum of 70% of the algal cells colonized just prior to bloom termination. Cross-correlation analyses between algal abundances and RCA bacterial colonization were statistically significant, in agreement with predator-prey models suggesting that RCA cluster bacteria caused algal bloom decline. Further investigation of molecular databases revealed that RCA cluster bacteria were numerically abundant during algal blooms sampled worldwide. Our findings suggest that the widespread RCA cluster bacteria may exert significant control over phytoplankton biomass and community structure in the oceans. We also suggest that coculture with phytoplankton may be a useful strategy to isolate and successfully grow previously uncultured but ecologically abundant marine heterotrophs.     

Protozooplankton in the Weddell Sea,Antarctica: Abundance and distribution in the ice-edge zone

Summary Protozooplankton were sampled in the iceedge zone of the Weddell Sea during the austral spring of 1983 and the austral autumn of 1986. Protozooplankton biomass was dominated by flagellates and ciliates. Other protozoa and micrometazoa contributed a relatively small fraction to the heterotrophic biomass. During both cruises protozoan biomass, chlorophyll a concentrations, phytoplankton production and bacterial biomass and production were low at ice covered stations. During the spring cruise, protozooplankton, phytoplankton, and bacterioplankton reached high concentrations in a welldeveloped ice edge bloom 100 km north of the receding ice edge. During the autumn cruise, the highest concentrations of biomass were in open water well-separated from the ice edge. Integrated protozoan biomass was <12% of the biomass of phytoplankton during the spring cruise and in the autumn the percentages at some stations were >20%. Bacterial biomass exceeded protozooplankton biomass at ice covered stations but in open water stations during the fall cruise, protozooplankton biomass reached twice that of bacteria in the upper 100m of the water column. The biomass of different protozoan groups was positively correlated with primary production, chlorophyll a concentrations and bacterial production and biomass, suggesting that the protozoan abundances were largely controlled by prey availability and productivity. Population grazing rates calculated from clearance rates in the literature indicated that protozooplankton were capable of consuming significant portions of the daily phyto- and bacterioplankton production.     

台湾海峡上升流区浮游植物对营养盐添加的响应

2006年6月在台湾海峡近岸上升流区通过表层水体营养盐添加的现场培养实验,研究该海区营养盐限制情况及其浮游植物水华产生的主要影响因素.对营养盐,叶绿素a浓度和浮游植物细胞丰度进行了测定,结果表明,实验中不存在明显的硅限制;氮磷营养盐均存在明显的限制,且氮限制情况更为严重.营养盐添加后,冰河拟星杆藻(Asterionellopsis glacialis)等硅藻迅速生长成为优势藻种,其对氮磷的利用机制有所不同.对氮营养盐采取吸收后迅速同化利用,相较于硝酸盐的补充,氨氮补充条件下优势硅藻更易迅速生长并迅速死亡;对磷营养盐的利用则由于体内磷库的存在,采用迅速吸收后贮存在体内慢慢消耗的利用机制.氮营养盐的补充是上升流期间浮游植物水华产生的主要因素.