Pliocene-Pleistocene alkenone, nitrogen and opal record of ODP Site 175-1082

In this study we present combined high-resolution records of sea surface temperature (SST), phytoplankton productivity, and nutrient cycling in the Benguela Upwelling System (BUS) for the past 3.5 Ma. The SST record provided evidence that upwelling activity off Namibia mainly intensified ca. 2.4-2.0 Ma ago in response to the cooling of the Southern Ocean and the resultant strengthening of trade winds. As revealed by productivity-related proxies, BUS intensification led to a major transition in regional biological productivity when considering the termination of the Matuyama Diatom Maximum (a diatom high-production event). Major oceanic reorganization in the Benguela was accompanied by nutrient source changes, as indicated by a new nitrogen isotopic (delta15N) record that revealed a stepwise increase at ca. 2.4 and ca. 1.5 Ma ago. The change in source region likely resulted from significant changes in intermediate water formation tied to the reorganization of oceanic conditions in the Southern Ocean, which may have in turn mainly controlled the global ocean N cycle, and therefore the N isotopic composition of nutrients since 3.5 Ma ago.

The nitrogen costs of photosynthesis in a diatom under current and future pCO2

With each cellular generation, oxygenic photoautotrophs must accumulate abundant protein complexes that mediate light capture, photosynthetic electron transport and carbon fixation. In addition to this net synthesis, oxygenic photoautotrophs must counter the light-dependent photoinactivation of Photosystem II (PSII), using metabolically expensive proteolysis, disassembly, resynthesis and re-assembly of protein subunits. We used growth rates, elemental analyses and protein quantitations to estimate the nitrogen (N) metabolism costs to both accumulate the photosynthetic system and to maintain PSII function in the diatom Thalassiosira pseudonana, growing at two pCO2 levels across a range of light levels. The photosynthetic system contains c. 15-25% of total cellular N. Under low growth light, N (re)cycling through PSII repair is only c. 1% of the cellular N assimilation rate. As growth light increases to inhibitory levels, N metabolite cycling through PSII repair increases to c. 14% of the cellular N assimilation rate. Cells growing under the assumed future 750 ppmv pCO2 show higher growth rates under optimal light, coinciding with a lowered N metabolic cost to maintain photosynthesis, but then suffer greater photoinhibition of growth under excess light, coincident with rising costs to maintain photosynthesis. We predict this quantitative trait response to light will vary across taxa.

(Table 1) Stable carbon and nitrogen isotopic ratios of potential prey species of bearded seals (Erignathus barbatus), Svalbard

Changing patterns of sea-ice distribution and extent have measurable effects on polar marine systems. Beyond the obvious impacts of key-habitat loss, it is unclear how such changes will influence ice-associated marine mammals in part because of the logistical difficulties of studying foraging behaviour or other aspects of the ecology of large, mobile animals at sea during the polar winter. This study investigated the diet of pregnant bearded seals (Erignathus barbatus) during three spring breeding periods (2005, 2006 and 2007) with markedly contrasting ice conditions in Svalbard using stable isotopes (d13C and d15N) measured in whiskers collected from their newborn pups. The d15N values in the whiskers of individual seals ranged from 11.95 to 17.45 per mil, spanning almost 2 full trophic levels. Some seals were clearly dietary specialists, despite the species being characterised overall as a generalist predator. This may buffer bearded seal populations from the changes in prey distributions lower in the marine food web which seems to accompany continued changes in temperature and ice cover. Comparisons with isotopic signatures of known prey, suggested that benthic gastropods and decapods were the most common prey. Bayesian isotopic mixing models indicated that diet varied considerably among years. In the year with most fast-ice (2005), the seals had the greatest proportion of pelagic fish and lowest benthic invertebrate content, and during the year with the least ice (2006), the seals ate more benthic invertebrates and less pelagic fish. This suggests that the seals fed further offshore in years with greater ice cover, but moved in to the fjords when ice-cover was minimal, giving them access to different types of prey. Long-term trends of sea ice decline, earlier ice melt, and increased water temperatures in the Arctic are likely to have ecosystem-wide effects, including impacts on the forage bases of pagophilic seals.

Biotic and abiotic parameters and cynobacteria abundance during an experimental set-up with varying pCO2 conditions

Heterocystous cyanobacteria of the genus Nodularia form extensive blooms in the Baltic Sea and contribute substantially to the total annual primary production. Moreover, they dispense a large fraction of new nitrogen to the ecosystem when inorganic nitrogen concentration in summer is low. Thus, it is of ecological importance to know how Nodularia will react to future environmental changes, in particular to increasing carbon dioxide (CO2) concentrations and what consequences there might arise for cycling of organic matter in the Baltic Sea. Here, we determined carbon (C) and dinitrogen (N2) fixation rates, growth, elemental stoichiometry of particulate organic matter and nitrogen turnover in batch cultures of the heterocystous cyanobacterium Nodularia spumigena under low (median 315 µatm), mid (median 353 µatm), and high (median 548 µatm) CO2 concentrations. Our results demonstrate an overall stimulating effect of rising pCO2 on C and N2 fixation, as well as on cell growth. An increase in pCO2 during incubation days 0 to 9 resulted in an elevation in growth rate by 84 ± 38% (low vs. high pCO2) and 40 ± 25% (mid vs. high pCO2), as well as in N2 fixation by 93 ± 35% and 38 ± 1%, respectively. C uptake rates showed high standard deviations within treatments and in between sampling days. Nevertheless, C fixation in the high pCO2 treatment was elevated compared to the other two treatments by 97% (high vs. low) and 44% (high vs. mid) at day 0 and day 3, but this effect diminished afterwards. Additionally, elevation in carbon to nitrogen and nitrogen to phosphorus ratios of the particulate biomass formed (POC : POP and PON : POP) was observed at high pCO2. Our findings suggest that rising pCO2 stimulates the growth of heterocystous diazotrophic cyanobacteria, in a similar way as reported for the non-heterocystous diazotroph Trichodesmium. Implications for biogeochemical cycling and food web dynamics, as well as ecological and socio-economical aspects in the Baltic Sea are discussed.

Physical, chemical and biological characteristics of disturbed and undisturbed Arctic lakes, Northwest Territories

1. Global warming is predicted to cause changes in permafrost cover and stability in the Arctic. Zones of high ion concentration in regions of ice-rich permafrost are a reservoir of chemicals that can potentially be transferred to fresh waters during thawing. Consequently, input of enriched runoff from the thaw and sediment and vegetation from the landscape could alter lakes by affecting their geochemistry and biological production. 2. Three undisturbed lakes and five lakes disturbed by retrogressive permafrost thaw slumps were sampled during late summer of 2006 to assess the potential effects of thermokarst shoreline slumping on water and sediment chemistry, the underwater light regime, and benthic macrophyte biomass and community structure. 3. Undisturbed lakes had sediments rich in organic material and selected micronutrients, while disturbed lakes had sediments richer in calcium, magnesium and strontium, greater transparency of the water column, and a well-developed submerged macrophyte community. 4. It is postulated that enriched runoff chemistry may alter nutrient availability at the sediment-water interface and also the degradation of organic material, thus affecting lake transparency and submerged macrophytes. The results suggest that retrogressive permafrost slumping can significantly affect food webs in arctic tundra lakes through an increase in macrophyte biomass and development of a more complex benthic habitat.

Concentrations of organic carbon, nitrogen and phosphorus on vertical profiles in waters of the tropical North Atlantic in March 2002

In the literature, an inconsistency exists between estimates of biotically-effected carbon export inferred from large-scale geochemical studies (Jenkins 1982; 47 gC m-2 a-1) and local measurements of turbulent nutrient supply (Lewis et al. 1986; 4 gC m-2 a-1) in the eastern subtropical North Atlantic. Nutrient supply to the upper ocean by turbulent mixing is reexamined using local standard oceanographic measurements and high-resolution vertical profiles of nutrients averaged over a large region directly comparable to that investigated by Jenkins (1982). Turbulent fluxes induced by internal waves and salt fingering, respectively, are separated according to Gregg (1989) and Zhang et al. (1998). Nutrient transport into the nutrient-consuming surface layer by salt fingering is more than fivefold higher than transport due to internal-wave induced turbulence. Still, this cannot resolve the above- mentioned apparent inconsistency, even if additional physical transport mechanisms such as eddy pumping, advection and horizontal diffusion are accounted for. Estimated nitrate fluxes due to vertical turbulent diffusion are 0.05-0.15 mol m-2 a-1, corresponding to 4-11 gC m-2 a-1. Observed NO3/PO4 turbulent flux ratios of up to 23 are interpreted as the imprint of N2 fixation.

(Table 1) Effects of desiccation on photochemical processes and nitrogenase activity in Nostoc commune s.l. colonies

The effects of desiccation on photochemical processes and nitrogenase activity were evaluated in Nostoc commune s.l. colonies in situ from a wet thufur meadow at Petuniabukta, Billefjorden, Central Svalbard, during the 2009 arctic summer. The colonies were collected in the fully hydrated state, and were subjected to slow desiccation at ambient temperatures (5 - 8°C) and low light (30 - 80 µmol/m**2/s). For each colony the weight, area, photochemical performance, and nitrogenase activity were determined at the beginning, as well as on every day during the first four days of the experiment; thereafter, on every second day until desiccation was complete. The photochemical performance was evaluated from variable chlorophyll fluorescence parameters (FV/FM, Phi(PSII) , qP, and NPQ), and the nitrogenase activity was estimated by an acetylene-ethylene reduction assay. A significant decrease in the photochemically active area was recorded from the third day, when the colony had lost approximately 40% of its original weight indicating some changes in the extracellular matrix, and stopped on the 14th to 18th day. No effects of the desiccation on the main photochemical parameters (FV/FM, Phi(PSII), qP) were observed up to the sixth to eighth days of desiccation. Slightly lower values of FV/FM and Phi(PSII) recorded in fully-hydrated colonies could be caused by impaired diffusion of CO2 into cells. The steep reduction of photochemical activity occurred between the eighth and tenth day of the experiment, when the colony had lost approximately 80% of its fully-hydrated weight. The nitrogenase activity was highest on the first day, probably due to improved diffusion of N2 into cells, then declined, but was detectable until the sixth day of the experiment. Since Nostoc commune s.l. colonies were capable of photosynthesis and nitrogen fixation to the level of ca. 60% of its fully-hydrated weight, even partly-hydrated colonies contribute substantially to carbon and nitrogen cycling in the High Arctic wet meadow tundra ecosystem.

Biological traits, and egg and fecal pellet production of Calanus finmarchicus and Calanus glacialis in situ and during experiments

The effects of temperature and food was examined for Calanus finmarchicus and C. glacialis during 3 phases of the phytoplankton spring bloom in Disko Bay, western Greenland. The 2 species were collected during pre-bloom, bloom, and post-bloom and exposed to temperatures from 0 to 10°C, combined with deficient or excess food. Fecal pellet and egg production were measured as indices for grazing and secondary production, respectively. Furthermore, changes in body carbon, nitrogen, and lipid content were measured. C. glacialis sampled before the bloom and incubated with excess food exhibited high specific egg production at temperatures between 0 and 2.5°C. Higher temperatures did not increase egg production considerably, whereas egg production for C. finmarchicus more than tripled between 2.5 and 5°C. Starved C. glacialis produced eggs at all temperatures stimulated by increasing temperatures, whereas starved C. finmarchicus needed temperatures above 5°C to produce eggs fueled by their lipid stores. Few C. finmarchicus had mature gonads at the initiation of the pre-bloom and bloom experiment, and egg production of C. finmarchicus therefore only increased as the ratio of individuals with mature gonads increased. During the bloom, both C. glacialis and C. finmarchicus used the high food availability for egg production, while refueling or exhausting their lipid stores, respectively. Finally, during the post-bloom experiment, production was low by C. finmarchicus, whereas C. glacialis had terminated production. Our results suggest that a future warmer ocean will reduce the advantage of early spawning by C. glacialis and that C. finmarchicus will become increasingly prevalent.