Epipelagic planktonic bioluminescence in the marginal ice zone of the Greenland Sea

Epipelagic planktonic bioluminescence in the marginal ice zone of the Fram Strait was investigated during April and May 1989. Vertical profiles of bioluminescence potential were measured using a HIDEX (High Input Defined EXcitation) design bathyphotometer. Mesozooplankton samples were collected with oblique tows using a 153 m mesh net. The amount of bioluminescence produced by mechanical stimulation of individual organisms was measured using an integrating sphere and photon-counting system. These measures of bioluminescence potential along with estimated abundances of bioluminescent organisms allow an independent estimated of integrated epipelagic bioluminescence potential. The zooplankton community structure was relatively simple in this region and the number of bioluminescent species correspondingly small. The majority of epipelagic bioluminescence was produced by copepods (Metridia spp.), larvaceans (Oikopleura spp.), euphausiids (Thysanoeassa spp.) and ostracods (Conchoecia spp.), although the relative contribution of these organisms to the overall bioluminescence potential varied considerably with time and location over the 2 mo period. Bioluminescent dinoflagellates were rare and did not contribute significantly to epipelagic bioluminescence. Integrated bioluminescence potential in the water column was significantly correlated with zooplankton biomass, but not with any other measured environmental parameter. No enhancement of chlorophyll, zooplankton biomass or bioluminescence was observed in the marginal ice zone compared to the adjacent open waters of the Fram Strait during this spring cruise.     

Contribution of salps to carbon flux of marginal ice zone of the Lazarev Sea, southern ocean

In order to estimate the in situ grazing rates of Salpa thompsoni and their implications for the development of phytoplankton blooms and for the sequestration of biogenic carbon in the high Antarctic, a repeat-grid survey and drogue study were carried out in the Lazarev Sea during austral summer of 1994/1995 (December/January). Exceptionally high grazing rates were measured for S. thompsoni at the onset of a phytoplankton bloom (0.2 to 0.8 μg chlorophyll a l⁻¹) in December 1994, with up to ≃160 μg of plant pigments consumed by an individual salp of 7 to 10 cm length per day. Dense salp swarms extended throughout the marginal ice zone, consuming up to 108% of daily phytoplankton production and 21% of the total chlorophyll a stock. Due to the much faster sinking rates and higher carbon content of salp faecal pellets, the efficiency of downward carbon flux through salps is much higher than through the other major grazers, krill and copepods. S. thompsoni can thus export large amounts of biogenic carbon from the euphotic zone to the deep ocean. With the observed ingestion rates during December 1994, this flux could have attained levels of up to 88 mg C m⁻² d⁻¹, accounting for the bulk of the vertical transport of carbon in the Lazarev Sea. However, in January 1995, when phytoplankton concentrations exceeded a threshold level of 1.0 to 1.5 μg chlorophyll a l⁻¹, salps experienced a drastic reduction in their feeding efficiency, possibly as a result of clogging of their filtering apparatus. This triggered a dramatic reversal in the relationship, during which a dense phytoplankton bloom developed in conjunction with the collapse of the salp population. Increases in the biomass and geographic range of the tunicate S. thompsoni have occurred in several areas of the southern ocean, often in parallel with a rise in sea-surface temperature during sub-decadal periods of warming anomalies. Received: 10 August 1997 / Accepted: 21 October 1997     

Large viruses and infected microeukaryotes in Ross Sea summer pack ice habitats

A variety of Ross Sea summer pack ice habitats between 66 and 75°S were examined for viruses 𔒦 nm capsid diameter. Maximum abundances of these viruses likely to infect eukaryotes were 10⁶-10⁷ ml^-1 brine in surface, interior, and bottom habitats and constituted up to 18% of the total (all sizes) viruses. There is abundant ultrastructural evidence for infection of a variety of microheterotrophs and some autotrophs. One station exhibited the classical characteristics of a lower latitude algal bloom with potential viral control. The blooming alga, Pyramimonas tychotreta Daugbjerg 2000, was infected, as were two abundant heterotrophs, Cryothecomonas spp. and an unidentified flagellate, that fed on P. tychotreta. Infections were observed in only one life history stage (multiflagellate cells) of P. tychotreta, suggesting a relationship among virus-induced lysis, life-history stages, physiology, and environmental factors regulating the life cycle. There is good evidence that diatoms are not a likely source of the large viruses, and viruses in general are not a major food source for ice microheterotrophs in summer.     

Effects of seasonal pack ice on the distribution of macrozooplankton and micronekton in the northwestern Weddell Sea

The presence of mesopelagic organisms in the guts of surface-foraging seabirds feeding in open areas within seasonal pack ice in the Antarctic has given rise to questions regarding the effects of pack ice on the underlying mesopelagic community. Bottom-moored free-vehicle acoustic instruments were used in concert with midwater trawls and baited traps to examine the abundance, size distribution and vertical distribution of pelagic organisms in the uppermost 100 m of the water column during the austral spring of 1992 in two areas of the northwestern Weddell Sea, one covered by seasonal pack icc and the other free of ice cover. Acoustic largets were more abundant and significantly larger at the open-water station than beneath pack ice. However, targets at the ice-covered site exhibited a pronounced diel pattern, with the largest targets detected only at night. Samples from night trawls at the icecovered site contained several species of large, vertically-migrating mesopelagic fishes, whereas these species were absent from trawls taken during the day. In addition, baited traps deployed in pack ice just beneath the ice-water interface collected large numbers of scavenging lysianassoid amphipods, while deeper traps beneath the ice and traps at the open-water station were empty, indicating the presence of a scavenging community associated with the undersurface of the ice. These results sapport the idea that mesopelagic organisms migrate closer to the surface beneath pack ice than in open water, exposing them to possible predation by surface-foraging seabirds.     

Under-ice amphipods in the Greenland Sea and Fram Strait (Arctic): environmental controls and seasonal patterns below the pack ice

During three "Polarstern" cruises to the ice-covered Greenland Sea (spring 1997, summer 1994, autumn 1995) studies on the under-ice habitat (morphology, hydrography, ice-algal biomass) and on the macrofaunal, autochthonous under-ice amphipods (species diversity, abundance) were carried out in order to describe environmental controls and seasonal patterns in this community. In spring, the ice underside was rather smooth and whitish, while in summer melting structures and sloughed-off ice-algal threads were observed, in autumn detritus clumps accumulated in depressions at the ice underside. Only in summer, a thin layer of warm (up to -0.6°C) and less saline (as low as S=6.3) water was found at the ice-water interface above Polar Water. Integrated ice-algal biomass was highest during autumn (2.6 mg chl a m^-2) and lowest during summer (1.2 mg chl a m^-2). Four species of under-ice amphipods occurred in spring and summer (Apherusa glacialis, Onisimus glacialis, O. nanseni, Gammarus wilkitzkii), but only the last species was observed at the ice underside in autumn. A. glacialis and G. wilkitzkii were equally abundant in spring; A. glacialis dominated in summer. The highest total abundance of amphipods occurred during summer (31.9 ind. m^-2), compared to lower abundances in spring and autumn (5.3 and 1.1 ind. m^-2, respectively). A factor analysis revealed seasonal patterns in the data set, which mainly influenced A. glacialis, and species-specific relations between several environmental factors and the distribution of under-ice amphipods. Abundance of A. glacialis was closely related to the under-ice hydrography and ice-algal biomass, whereas the other amphipod species were more influenced by the under-ice morphology. It is therefore stated that the observed thinning of the Arctic sea ice and the resulting increased meltwater input and change in morphology of floes will have a profoundly adverse effect on the under-ice amphipods.     

Diel patterns of planktonic bioluminescence in the northern Sargasso Sea

Day-night changes in the vertical distribution, intensity, and size of bioluminescence flashes were investigated during a series of cruises to the northern Sargasso Sea in 1987 and 1988. Overall, depth integrated bioluminescence potential and flash density estimated from in situ measurements with a pumping bathyphotometer were 2 to 5 x higher at midnight than at midday. Depths from 50 to 100 m exhibited the most substantial day to night increases in bioluminescence potential and flash density. When classified by flash size (photon output per flash event), the increase from day to night was significant for all flash sizes, but was most dramatic for small flashes producing <7 x 10⁸ photons flash^-1. Bioluminescence potential and flash density increased 2 to 3 x during bathyphotometer measurements made at dusk. Bioluminescent light budgets estimated from day and night net collections in May and August 1987 also predicted 2.5 x higher nighttime than daytime mesoplankton bioluminescence. However, large bioluminescent taxa (mesoplankton) capable of significant vertical migrations only contributed on the order of 15% of the total bioluminescence in surface waters. Our results do not support the idea that most of the nightly increase in bioluminescence potential and flash density are due to vertical migration of bioluminescent organisms; rather they are consistent with an alternate view that photoinhibition of bioluminescent flashing by dinoflagellates may be primarily responsible for the diel patterns.     

Observations of planktonic bioluminescence in the euphotic zone of the California Current

The distributions of bioluminescence, temperature, salinity, oxygen. pH, and chlorophyll a were measured at 10 m intervals, to a depth of 100 m at a station (33°46N; 119°36W) in the California Current from 17 to 20 July 1982. The distribution of bioluminescence showed a marked day-night change which was consistent over the sampling period. The nighttime maximum was at the surface, and the daytime maximum was between 30 and 40 m. The shapes of the day and night distributions were independent of the absolute intensity of bioluminescence and were also insensitive to advection, as inferred from changing temperature-salinity relationships. The nighttime depth distribution broadened during a period of high wind Day to night differences in the color spectrum at the depth of maximum bioluminescence suggest that the luminescent organisms differed from day to night.     

Vertical fine structure of the biomass and composition of algal communities in Arctic pack ice

The biomass and composition of algal communities in sea ice were studied during two summer expeditions to the central Arctic Ocean and the Greenland Sea. Based on algal pigment determination and cell counts, high biomass accumulations were found at the surface, in the interior and in the bottom layer of the ice floes. Pennate diatoms dominated in the bottom layer, while phototrophic flagellates and cysts of unknown origin were the most abundant taxa in the upper parts. The lowermost 20 to 40 cm contained between 4 and 62% of the entire algal biomass. Consequently, ice biological studies, which deal only with the bottom few centimetres of the ice floes, will underestimate algal biomass and production by factors of up to 25. Differences between the results of this study and published data from coastal locations point towards different biological regimes in Arctic sea ice. The algal biomass in coastal ice is about two orders of magnitude higher and composed mainly of diatoms, probably supported by nutrient influx from the water column. In the pack ice of the central Arctic, nutrient supply is probably reduced, and flagellates contribute substantially to total algal biomass. However, methodological problems might partially be responsible for the observed differences. Received: 12 June 1998 / Accepted: 11 December 1998     

Seasonal variations in distribution patterns of sympagic meiofauna in Arctic pack ice

During two expeditions of the R.V. Polarstern to the Arctic Ocean, pack ice and under-ice water samples were collected during two different seasons: late summer (September 2002) and late winter (March/April 2003). Physical and biological properties of the ice were investigated to explain seasonal differences in species composition, abundance and distribution patterns of sympagic meiofauna (in this case: heterotrophs >20 µm). In winter, the ice near the surface was characterized by extreme physical conditions (minimum ice temperature: –22°C, maximum brine salinity: 223, brine volume: 5%) and more moderate conditions in summer (minimum ice temperature: –5.6°C, maximum brine salinity: 94, most brine volumes: 5%). Conditions in the lowermost part of the ice did not differ to a high degree between summer and winter. Chlorophyll a concentrations (chl a) showed significant differences between summer and winter: during winter, concentrations were mostly <1.0 µg chl a l^–1, while chl a concentrations of up to 67.4 µmol l^–1 were measured during summer. The median of depth-integrated chl a concentration in summer was significantly higher than in winter. Integrated abundances of sympagic meiofauna were within the same range for both seasons and varied between 0.6 and 34.1×10³ organisms m^–2 in summer and between 3.7 and 24.8×10³ organisms m^–2 in winter. With regard to species composition, a comparison between the two seasons showed distinct differences: while copepods (42.7%) and rotifers (33.4%) were the most abundant sea-ice meiofaunal taxa during summer, copepod nauplii dominated the community, comprising 92.9% of the fauna, in winter. Low species abundances were found in the under-ice water, indicating that overwintering of the other sympagic organisms did not take place there, either. Therefore, their survival strategy over the polar winter remains unclear.Communicated by O. Kinne, Oldendorf/Luhe     

Phytoplankton in the Weddell Sea,Antarctica: composition,abundance and distribution in water-column assemblages of the marginal ice-edge zone during austral autumn

At the compacted, north-south line of the ice edge, phytoplankton were sampled during early austral autumn of 1986 in the northwestern Weddell Sea. Cells from discrete water bottle samples from 12 stations on two east-west transects were counted to gain quantitative information on the composition, abundance, distribution, and condition of the phytoplankton in water-column assemblages. Over 70 species were found. The highest numbers of total cells (integrated through the top 150 m) were found in open water, well-separated from and to the east of the ice edge on the southern transect, with 6.01×10¹⁰ cells m^-2. The relative abundance of diatoms was low at ice-convered stations (< 35% of the total phytoplankton in preserved samples) and high at open-water stations (> 80%); however, the relative abundance of the prymnesiophyte Phaeocystis sp. was high at ice-covered stations (> 60%) and low at open-water stations (< 16%), with lower absolute abundances than during a previous austral-spring phytoplankton increase. In the open ocean, the dominants were the pennate diatoms Fragilariopsis cylindrus, Pseudonitzschia prolongatoides, F. curta, and a small form of the centric diatom Chaetoceros dichaeta in chains. Although the three pennate diatoms were frequently dominant in number, they represented less biomass than C. dichaeta in open waters. Mean phytoplankton abundance was low (0.2×10⁶ cells l^-1) but, overall, the diatom cell density (0.14×10⁶ cells l^-1) was similar to that found previously during a northward transect from ice-covered to ice-free water at the Weddell-Scotia Sea ice edge (spring 1983). The phytoplankton spatial patterns in the two autumn transects differed, with the more southerly transect exhibiting a higher abundance of diatoms and dinoflagellates. The ratio of full to empty diatoms was higher on the southern transect, indicating a healthy population, while lower ratios of full/empty frustules on the northern transect suggested a generally declining population. However, Phaeocystis sp. was more abundant on the northern transect.     

Pattern of planktonic bioluminescence in the northern Sargasso Sea: Seasonal and vertical distribution

The vertical structure of bioluminescence potential (BPOT) and flash density (FD) were measured on five cruises to the northern Sargasso Sea in 1987 and 1988. Depth-integrated (0 to 150 m) BPOT did not vary seasonally, remaining within the range 9 to 15 × 10¹⁵ photons m^–2 in all months sampled. Conversely, depth-integrated FD was significantly higher (> 2 × 10⁵ flashes m^–2) during winter (November and March) than during summer (< 9 × 10⁴ flashes m^–2 in May and August). The vertical patterns of BPOT and FD were well correlated within a single profile, more highly so in summer than in winter. Despite intracruise variability in the vertical pattern of BPOT and FD, there were clear summer-winter differences in the vertical distribution of BPOT and FD. During winter, BPOT and FD were maximal and relatively uniform throughout the surface mixed layer; for example in November they declined sharply within the thermocline at 130 to 150 m. During summer, BPOT and FD were greatest (12 to 25 × 10¹³ photons m^–3 and 600 to 1 200 flashes m^–3, respectively) at subsurface depths. Commonly in summer, the upper depth limit of high BPOT and FD occurred at the base of the surface mixed layer (10 to 40 m) and the lower depth limit was located at the base of the subsurface fluorescence maximum (usually at 100 to 120 m).     

Spectral composition of bioluminescence of epipelagic organisms from the Sargasso Sea

The spectral characteristics of single identified epipelagic sources of bioluminescence from the western Sargasso Sea were measured with an optical multichannel analyzer (OMA) system during the April, 1985, Biowatt cruise. The emission spectra of specimens representing 45 species from 8 phyla were measured. Peak bioluminescence emissions typically occurred between 440 and 500 nm, in the blue region of the visible spectrum. Three exceptions involved emission in the green, yellow, and red spectral regions. Intraspectific variability in spectra, was noted in several species. One shrimp species exhibited two modes of light emission, each with different emission spectra. Other cases involved dynamic color shifts of 10 to 14 nm; the source of the spectral variability is unknown, but may involve optical filtering or differences in the color of luminescence from multiple sites of light emission. Measurements from independent samples of unsorted plankton revealed different spectral distributions. This suggests that the spectral emissions of bioluminescence in the upper water column will vary, based on species assemblage.     

Sea ice retreat alters the biogeography of the Bering Sea continental shelf.

Seasonal ice cover creates a pool of cold bottom water on the eastern Bering Sea continental shelf each winter. The southern edge of this cold pool, which defines the ecotone between arctic and subarctic communities, has retreated approximately 230 km northward since the early 1980s. Bottom trawl surveys of fish and invertebrates in the southeastern Bering Sea (1982-2006) show a coincident reorganization in community composition by latitude. Survey catches show community-wide northward distribution shifts, and the area formerly covered by the cold pool has seen increases in total biomass, species richness, and average trophic level as subarctic fauna have colonized newly favorable habitats. Warming climate has immediate management implications, as 57% of variability in commercial snow crab (Chionoecetes opilio) catch is explained by winter sea ice extent. Several measures of community distribution and structure show linear relationships with bottom temperature, suggesting warming climate as the primary cause of changing biogeography. However, residual variability in distribution not explained by climate shows a strong temporal trend, suggesting that internal community dynamics also contribute to changing biogeography. Variability among taxa in their response to temperature was not explained by commercial status or life history traits, suggesting that species-specific responses to future warming will be difficult to predict.     

Distribution of coccolithophores in marginal seas along the western Pacific Ocean and in the Red Sea

The distribution of coccolithophores was studied in the neritic environment along the western margin of the Pacific Ocean: the Inland Sea of Seto, Yellow Sea, East China Sea, South China Sea, Java Sea, Timor Sea, Arafura Sea and Gulf of Carpentaria. The coccolithophore community in the Red Sea was also studied for comparison with the Pacific marginal seas. With minor exceptions, the coccolithophore communities were very similar throughout the neritic areas investigated, but differed completely from the pelagic community in three aspects. Firstly, almost all neritic coccolithophores, regardless of species, suffered various degrees and forms of malformation with relation to the morphology of their coccoliths, while such malformation was rare in the pelagic population. Nitrogen deficiency may cause such malformation. Secondly, the diversity of species in these marginal seas was much lower than in the pelagic environment, although no species was found to be exclusively neritic. Emiliania huxleyi, usually ubiquitous in oceanic areas and in various neritic environments of higher latitudes, was scarce, while Gephyrocapsa oceanica dominated the flora throughout the studied areas. Finally, the horizontal and vertical distributions of the neritic populations were sporadic compared to those of the rather uniform pelagic environments.     

Total labile carbon in the euphotic zone of the Baltic Sea as measured by BOD

Total labile organic carbon was estimated in the euphotic zone of the Baltic Sea by a method based on the determination of the biological oxygen demand (BOD); the method is extremely simple, requires little sample water, and is quite precise (±4.5 μmol O₂/l or ±40.4 μgC_lab/l). The amount found was 1,150 μgC_lab/l, or about 29% of the total organic carbon. At 1 m, a relationship between biological activity, as measured by chlorophyll content, and the amount of labile carbon was found: ln C_lab (μg/l)=ln 6.87 (μg/l) +0.272 ln chlorophyll (μg/l), with a correlation coefficient of 0.86, which is significantly greater than zero at the 0.001 level.     

Estimating carbon flux to pelagic grazers in the ice-edge zone of the eastern Bering Sea

Zooplankton ingestion of phytoplankton carbon in the iceedge zone of the Eastern Bering Sea was measured using a deck incubation approach in 1982. Using further samples collected in 1983, the plant cell carbon to cell volume ratio was estimated at 0.0604 pg m^–3 from an experimentally determined particulate carbon to seston volume relationship. The application of this conversion to the results of experimental incubations of natural plant stocks with net-caught zooplankton produced ingestion rates of 68.8 and 10.26 mg C g^–1 grazer d^–1 for copepods and euphausiids, respectively. Extrapolating these rates to in situ zooplankton biomass at the edge of the seasonal ice pack yielded carbon flux rates through the zooplankton community ranging between 6.5 and 32.8 mg C m^–2 d^–1. This consumption amounted to less than 2% of the daily phytoplankton production in the ice-edge zone.     

Sea ice microbial production supports Ross Sea benthic communities: influence of a small but stable subsidy

Diversity in guilds of primary producers enhances temporal stability in provision of organic matter to consumers. In the Antarctic ecosystem, where temporal variability in phytoplankton production is high, sea ice contains a diatom and microbial community (SIMCO) that represents a pool of organic matter that is seasonally more consistent, although of relatively small magnitude. The fate of organic material produced by SIMCO in Antarctica is largely unknown but may represent an important link between sea ice dynamics and secondary production in nearshore food webs. We used whole tissue and compound-specific stable isotope analysis of consumers to test whether the sea ice microbial community is an important source of organic matter supporting nearshore communities in the Ross Sea. We found distinct gradients in delta13C and delta15N of SIMCO corresponding to differences in inorganic carbon and nitrogen acquisition among sites with different sea ice extent and persistence. Mass balance analysis of a suite of consumers demonstrated large fluxes of SIMCO into the nearshore food web, ranging from 5% to 100% of organic matter supplied to benthic species, and 0-10% of organic matter to upper water column or pelagic inhabitants. A delta13C analysis of nine fatty acids including two key biomarkers for diatoms, eicosapentaenoic acid (EPA, 20:5omega3), and docosahexaenoic acid (DHA, 22:6omega3), confirmed these patterns. We observed clear patterns in delta13C of fatty acids that are enriched in 13C for species that acquire a large fraction of their nutrition from SIMCO. These data demonstrate the key role of SIMCO in ecosystem functioning in Antarctica and strong linkages between sea ice extent and nearshore secondary productivity. While SIMCO provides a stabilizing subsidy of organic matter, changes to sea ice coverage associated with climate change would directly affect secondary production and stability of benthic food webs in Antarctica.     

Localization of bioluminescence in the siphonophore Nanomia cara

The base of the tentacle of the developing physonect larva (Nanomia cara) has a bioluminescent region. The ability to produce light in the larva is transitory; this ability first appears at about two days of development and is disappearing by eight days, as the larva begins to feed. Subsequently paired bilaterally symmetrical bioluminescent organs are found on the nectophores and the bracts of the adult colony. In both the larva and the adult, bioluminescence is mediated by a calcium specific photoprotein. In all cases the photocytes lack a green fluorescent protein.