ALBERT

Description: A predictive model of carbon isotope fractionation (sigma p) and abundance (delta13C phyto) is presented under circumstances where photosynthesis is strictly based on CO2(aq) that passively diffuses into marine phytoplankton cells. Similar to other recent models, the one presented here is based on a formulation where the expression of intracellular enzymatic isotope fractionation relative to that imposed by CO2(aq) transport is scaled by the ratio of intracellular to external [CO2(aq)], ci/ce. Unlike previous models, an explicit calculation of ci is made that is dependent on ce as well as cell radius, cell growth rate, cell membrane permeability to CO2(aq), temperature, and, to a limited extent, pH and salinity. This allows direct scaling of ci/ce to each of these factors, and thus a direct prediction of sigma p and delta13C phyto responses to changes in each of these variables. These responses are described, and, where possible, compared to recent experimental and previous modeling results.

Description: The conventional model whereby plume volcanism forms linear age-progressive volcanic chains, with the youngest activity occurring nearest a spreading axis (at a 'hotspot'), has been challenged for the Easter seamount chain1–4. Whereas early work suggested the existence of a linear melting anomaly (a 'hotline')1,2, more recent studies3,4 have proposed a hotspot near Salas y Gomez island, connected with the Easter microplate spreading system by an ~800-km-long, volcanically active plume channel. Here we use geochemical, geological and geochronological data to argue that the hotspot lies close to Easter Island. Moreover, new isotopic data for lavas from the seamount chain provide evidence for bidirectional flow between the spreading axis and the plume, thus supporting geophysical and fluid-dynamical models of mantle flow in a plume/spreading axis system5–7. Material balance and flux considerations show the Easter plume to be weak and cool compared with those beneath larger features such as Iceland, Hawaii and the Galápagos islands.

Description: A knowledge of past changes in the biological productivity of the oceans is important for understanding the interactions between carbon cycling and climate. Phytoplankton productivity in today's oceans can be estimated from the concentrations of chlorophyll in sea water1, but chlorophyll is not preserved in the sediments. Existing proxies for past algal productivity do not represent total productivity; for example, biogenic opal2 reflects the contribution of only part of the phytoplankton community, and the organic carbon record can be subject to contamination from terrestrial inputs2,3. Although chlorins, the pigment-transformation products of chlorophyll, are widespread in Quaternary marine sediments, their potential as proxy measures of past variations in primary productivity has not been convincingly demonstrated. Here we report a high-resolution molecular stratigraphic record of chlorin concentrations over the past 350,000 years in a sediment core from the subtropical Atlantic continental margin. Maxima in the chlorin accumulation rate coincide with significant peaks in the accumulation rates of biogenic opal (at the end of glacial terminations) and organic carbon (between terminations). These results suggest that chlorins, unlike other proxies, can serve as a measure of total primary productivity variations.

Description: Nutrient competition experiments were performed with periphytic microalgae from the Kiel Fjord, western Baltic Sea. Diatoms were dominant competitors at high Si:N ratios (under N-limited conditions) and high Si:P ratios (under P-limited conditions). Under low silicate supply, cyanobacteria became dominant at low N:P ratios and chlorophytes at high ones. Changes in light intensity influenced the outcome of competition at the species level but not at the level of higher taxa. The nitrogen source (ammonium or nitrate) had no impact on the outcome of competition.

Description: Cod larvae from laboratory rearing experiments aged from 1 to 12 d after hatching, both fed and deprived of food, were analysed. The number of increments on the otolith and the width of these increments were determined together with the RNA/DNA ratios on the same individual larva. Alizarin marking of the otoliths was performed to confirm the formation of daily increments. Cod larvae reared at 6*C formed the first ring right after hatching and deposited increments on a daily basis. A comparison of the measurements between the right and the left lapillus showed that these can differ, if the radius is taken. The lapillae core showed especially high individual variability, whereas the sum of the increments did not differ between both lapilli. Until Day 10 after hatching, while the larvae were still feeding on their yolk, the external food situation did not affect the increment width of the lapilli or the RNA/DNA ratios. In larvae older than 10 d the width of the daily increments was dependent on the nutritional situation and RNA/DNA ratios decreased in starving larvae in comparison to feeding larvae. RNA/DNA ratios and increment widths were correlated.

Description: Mixed cultures of 4 polar diatoms regularly found in Antarctic pack ice were grown over 20 d in closed bottles at high light (200 to 250 umol photons m-2 s-1) and at 0*C in order to investigate growth physiology and biomass production under conditions simulating the sea ice habitat during summer. Species tested were: Chaetoceros cf. neogracile, Fragilariopsiscylindrus, Thalassiosiraantarctica and Porosira pseudodenticulata. Initially, all species grew exponentially, but exponential growth ceased for P. pseudodenticulata and T.antarctica after 6 d, for F.cylindrus after 8 d, and for C. cf. neogracile after 10 d. Slight increases in cell number were observed for all species 2 d later. Peak biomass amounted to 140 ug chl a (850 umol particulate organic carbon, POC) l-1. At the same time, concentrations of dissolved inorganic carbon (DIC) were reduced by 1000 uM, oxygen concentrations increased to 1400 uM, and pH increased to 10.5. At this stage, a substantial decline in plasma-containing cells was recorded for F.cylindrus. C. cf. neogracile accounted for 80%, and C. cf. neogracile and F.cylindrus accounted for 〉95% of total carbon biomass. The carbon isotope composition of POC (expressed as delta13C) increased from -24 to -9 during the experiment. Model calculations showed that diffusive uptake of dissolved CO2 satisfied cellular carbon demand for all species except P. pseudodenticulata at CO2(aq) concentrations 〉0.5 uM, whereas direct HCO3- utilization was observed for C. cf. neogracile below this concentration. Our data confirm that intense photosynthetic carbon assimilation may lead to profound chemical changes in isolated interstitial brine solutions, with significant consequences for sea ice biota. We propose that the capacity to efficiently utilize ambient DIC, possibly mediated by virtue of favorable surface to volume ratios as well as active pathways of inorganic carbon acquisition, favors growth of small diatoms, and may be an important factor driving ice algal species succession during summer blooms. Since only 2 species continued to grow in fresh medium following experimental incubation (C. cf. neogracile and P. pseudodenticulata), differential tolerance to chemical variations may influence the seeding potential of ice algae following release into the open water.

Description: A fundamental issue in marine science is the identification of the factors controlling biological uptake of CO2, in high-nitrate, low-chlorophyll regions. A recent in situ iron fertilization experiment demonstrated that iron limitation is responsible for low phytoplankton stocks in the equatorial Pacific4. Here we show that flavodoxin, a biochemical marker of iron limitation, can be used to map the degree of iron stress in natural populations. Flavodoxin assays along a 900-km east-west transect in the northeastern subarctic Pacific revealed a pronounced increase in iron stress in the region west of the 135° W meridian. Addition of dissolved iron alleviated this stress. Immunostaining of single cells from the most western station showed that flavodoxin is present specifically within the chloroplasts of diatoms. Our approach provides a rapid means of defining the extent of iron stress in the ocean5 and supports the hypothesis that diatoms are iron stressed in the northeast Pacific.

Description: Epibiosis is one of the closest interspecies associations. The presence of epibionts potentially causes a multitude of beneficial or detrimental effects for the basibiont. It has been shown previously that large epibionts may increase the risk of dislodgement of bivalves. In this study, sublethal effects of epibiont-induced drag increase are investigated. I assessed (1) the effects of common epibiont species (Balanus improvisus, Enteromorpha intestinalis, Ectocarpus sp.) on drag properties of the host (the periwinkle Littorina littorea), and (2) the long-term consequences of drag increase on growth rates of snails living in steady flow. All epibiont species increase drag on the host snail. They do so to unequal extents. This may be due to morphological and hydrodynamic differences among the epibionts. Thus, per unit volume of epibiont, the filamentous alga Ectocarpus sp, has a substantially stronger effect than the barnacles. Synergistic effects on drag increase can be observed in a mixed aufwuchs community. As compared to clean conspecifics, snails bearing artificial epibionts grow 35% more slowly when exposed to moderate, steady flow (8 cm s(-1)) for 5 mo. This difference in growth rates is enhanced when food is limited. I hypothesize that fouled snails coping with higher drag invest more energy into foot activities (muscles and mucus). As a consequence, when food is limited, growth rates decrease in fouled snails.

Description: The dynamics of methane production and oxidation in the sediments of Kiel Harbour (Baltic Sea) were studied over a period of 2 yr. Experimentally determined rates of methanogenesis varied between 13 and 82 mu mol CH4 m(-2) h(-1) integrated over the top 30 cm of sediment. The zone of maximum production was between 20 and 30 cm depth. High potential rates could be induced in sulfate-containing surface layers by the addition of molybdate, acetate or methanol, indicating competition with sulfate-reducing bacteria. The methane content in the sediment pore water increased from the surface to a seasonally varying depth of between 5 and 20 cm, where sulfate concentrations approached zero and methane concentrations of 3 to 4 mM were recorded. Aerobic methane oxidation rates in the surface sediment layer were in most cases higher than the methane flux expected to reach the sediment surface. Oxygen-depleted or anoxic conditions in the deep water, which frequently occurred during stratification periods in summer, resulted in methane gradients with increasing concentrations from the chemocline to the sediment. After recirculation of the water column in autumn in both years, unexpectedly high methane concentrations were also measured in the oxic water column of Kiel Harbour. This coincided with very reduced conditions in the surface sediment that allowed measurable methane production in the 0 to 10 cm layer.

Description: It has often been suggested that, given their large food requirements, sperm whale Physeter macrocephalus distributions should reflect the distribution of productive ocean environments, and it seems therefore that chlorophyll concentration might be a good indicator of sperm whale distribution. To examine the existence of such a relationship, and to determine over which sclaes it occurs, sperm whale density was correlated with phytoplankton pigment concentration over a range of spatial and temporal scales. Sperm whale distribution was detrmined using records of kills from 19th century Yankee whaling, and the distribution of pigment concentration from satellite colour observations averaged over 8yr interval. These measures were compared over scales of 220km square to 1780km square. The distribution of sperm whales in the temperature and tropical Pacific Ocean was associated with distributions of phytoplankton pigment over every spatial scale considered, and the coefficient of correlation increased with increasing spatial scale. However, a few exceptions to this scheme were found, implying that other factors would be of importance in some regions. This study confirmed the existence of space lag and a time lag between a peak in chlorophyll concentration and a peak in sperm whale density. It also demonstrated that over large spatial scales, and when the data are averaged over large temporal scales, chlorophyll concentration is a good indicator of sperm whale distribution and that over these scales ocean colour recorded from space could help predict areas of high or low sperm whale density.

Description: Predator data and exploratory fishing in the Scotia Sea have revealed the presence of cephalopod stocks in the Antarctic Polar Frontal Zone (PFZ). This is a vast, remote region where large epipelagic cephalopods aggregate into highly mobile schools making them difficult to locate and sample. We used satellite tagged predators and shipboard acoustics for coarse and fine scale location of cephalopods concentrations, and sampled them with commercial and scientific nets to determine the relationship between cephalopod distribution and mesoscale oceanographic features at the PFZ. Saltellite tags were attached to 9 grey-headed albatrosses Diomedea chrysostoma, breeding at Bird Island, South Georgia, to monitor foraging at sea in January-March 1994. A foraging area at the PFZ, north of South Georgia, was located, an acoustic survey undertaken and a fixed situation established where gular midwater trawl 25m² (RTM25), a horizontal multiple plankton sampler and a neuston net. Acoustic layers were targeted and the RMT25 sampled 200m layers to 1000m in daylight and darkness. Cephalopods were simultaneously recovered from food samples fed to D. chrysostoma chicks at Bird Island. Two CTD transects, approximately normal to the major current flow, were undertaken across the PFZ and remote-sensed-sea-surface temperature images from NOAA polar orbiting satellites that exploited by D. chrysostoma. The largest and most conspicious species was the ommastrphid squid Martialia hyadesi which is the most important cephalopod prey species. Net-sampled M. hyadesi had been feeding on crustaceans and mesopelagic fish. The cephalopod community was sampled in a feature, interpreted as a warm core ring, in an area characterised by mesoscale features associated with the bathymetry of the northern end of the Northeast Georgia Rise and near a gap in the Falkland Ridge. The association of these mesoscale features with the bathymetry suggests that they may be predictable foraging locations for the cephalopods and their predators.

Description: The accuracy of species-specific phytoplankton growth rates estimated by cell cycle analysis was tested with the dinoflagellate Prorocentrum minimum (Pav.) Sch. under conditions of altered nitrogen and phosphorus availability. Reduced nutrient availability caused major changes in the duration of cell cycle phases. At the nutrient level of complete f/2 media, the length of the combination of S, G2, and M phases was about 8 h at growth rates of 0.53 to 0.56 d-' A decrease in ~ 0 ,o~r N-O3 concentration extended the S+G2+M phase to about 15.5 to 17.7 h at growth rates ranging from 0.41 to 0.30 d-' Changes in phase durations dld not significantly affect growth rate estimates. In addition, a minimum growth rate, calculated from the maximum values on phase fraction curves, was shown to be usable as an error detector in some cases. Results support the validity of cell cycle analysis to measure in situ growth rates.

Description: ISOTOPE ratios and concentrations of incompatible trace elements are remarkably successful in discriminating the tectonic origin and magmatic source components for basalts1–5. But problems remain with discriminating the tectonic origin of some tholeiites, especially where field relations and other geological evidence are ambiguous. For example, the tectonic origin of basalts from the Troodos ophiolite (Cyprus) has been debated for several decades. Most workers have been unable to distinguish between an island-arc and/or back-arc origin for the ophiolite6–8. Here we use volatile, K2O and TiO2 contents from ∼250 fresh submarine volcanic glasses to discriminate between tholeiites from different tectonic regimes. K2O÷H2O ratios are lower (〈0.70) in spread ing-centre glasses than in those from island arcs and intraplate oceanic islands. Back-arc-basin basalts can generally be separated from mid-ocean-ridge basalts by their high H2O contents. Using this information, we show that some fresh glasses from the Troodos ophiolite have a clear back-arc-basin affinity.