Tab.1: Total estimated muskox numbers and quotas in management zones of the Northwest Territories and Nunavut

In view of the drastic growth in the Canadian Inuit population, the rising costs of living, the missing job and income alternatives and the high unemployment rate in the arctic, efforts are being made to make use of the muskox populations in order to provide additional sources of food and/or revenue. The present paper attempts to review the course of muskox utilization in the Canadian Arctic and to tentatively assess its present as weIl as its future economic importance. Starting with the pre-European status of muskoxen in Canada, the drastic reduction in numbers resulting from the combined efforts of hide traders, whalers and expedition parties in the 19th and early 20th centuries, the impact of the legal protection and the recovery since 1917 are being described. Establishing muskox farms with semi-domesticated herds failed in Canada in the 1970's. Since 1969, though, increasing numbers of animals have been allotted to many Inuit communities, and despite the fact that most of the animals were primarily used for subsistence purposes, some communities could reserve part of their quotas for trophy (sport) hunters. While controlled sustainable subsistence and trophy hunts may eventually be carried out over the whole muskox range, including recently colonized northern Quebec, commercial harvesting for meat, hides and wool, introduced in 1981, will at least for some time be restricted to Banks and Victoria islands which at present show 78 % of the Canadian muskox population and 94 % of the overall quota.

Growth rates and biometric measurements of coccolithophores (Coccolithus pelagicus, Coccolithus braarudii, Emiliania huxleyi) during experiments

Coccolithophores, a diverse group of phytoplankton, make important contributions to pelagic calcite production and export, yet the comparative biogeochemical role of species other than the ubiquitous Emiliania huxleyi is poorly understood. Here we examined the relative importance of E. huxleyi and two Coccolithus species (Coccolithus pelagicus and Coccolithus braarudii), in terms of daily calcite production, by culturing E. huxleyi and Coccolithus in parallel, and comparing growth rates and biometrically determined cellular carbon calcite quotas. Biometric measurements of Coccolithus species, and E. huxleyi cell diameters, were performed using polarised light microscopy. Scanning electron microscopy was used for all other biometric measurements of E. huxleyi.

Ocean acidification decreases the light-use efficiency in an Antarctic diatom under dynamic but not constant light, link to supplementary data

There is increasing evidence that different light intensities strongly modulate the effects of ocean acidification (OA) on marine phytoplankton. The aim of the present study was to investigate interactive effects of OA and dynamic light, mimicking natural mixing regimes. The Antarctic diatom Chaetoceros debilis was grown under two pCO2 (390 and 1000 latm) and light conditions (constant and dynamic), the latter yielding the same integrated irradiance over the day. To characterize interactive effects between treatments, growth, elemental composition, primary production and photophysiology were investigated. Dynamic light reduced growth and strongly altered the effects of OA on primary production, being unaffected by elevated pCO2 under constant light, yet significantly reduced under dynamic light. Interactive effects between OA and light were also observed for Chl production and particulate organic carbon (POC) quotas. Response patterns can be explained by changes in the cellular energetic balance. While the energy transfer efficiency from photochemistry to biomass production (Phi_e,C) was not affected by OA under constant light, it was drastically reduced under dynamic light. Contrasting responses under different light conditions need to be considered when making predictions regarding a more stratified and acidified future ocean.

Seawater carbonate chemistry and processes during experiments with cyanobacterium Trichodesmium (IMS101), 2009

We investigated carbon acquisition by the N2-fixing cyanobacterium Trichodesmium IMS101 in response to CO2 levels of 15.1, 37.5, and 101.3 Pa (equivalent to 150, 370, and 1000 ppm). In these acclimations, growth rates as well as cellular C and N contents were measured. In vivo activities of carbonic anhydrase (CA), photosynthetic O2 evolution, and CO2 and HCO3- fluxes were measured using membrane inlet mass spectrometry and the 14C disequilibrium technique. While no differences in growth rates were observed, elevated CO2 levels caused higher C and N quotas and stimulated photosynthesis and N2 fixation. Minimal extracellular CA (eCA) activity was observed, indicating a minor role in carbon acquisition. Rates of CO2 uptake were small relative to total inorganic carbon (Ci) fixation, whereas HCO{3 contributed more than 90% and varied only slightly over the light period and between CO2 treatments. The low eCA activity and preference for HCO3- were verified by the 14C disequilibrium technique. Regarding apparent affinities, half-saturation concentrations (K1/2) for photosynthetic O2 evolution and HCO3- uptake changed markedly over the day and with CO2 concentration. Leakage (CO2 efflux : Ci uptake) showed pronounced diurnal changes. Our findings do not support a direct CO2 effect on the carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase (RubisCO) but point to a shift in resource allocation among photosynthesis, carbon acquisition, and N2 fixation under elevated CO2 levels. The observed increase in photosynthesis and N2fixation could have potential biogeochemical implications, as it may stimulate productivity in N-limited oligotrophic regions and thus provide a negative feedback in rising atmospheric CO2 levels.

Polar membrane lipids and elemental composition of Thermococcus kodakarensis cells according to growth stage and phosphorus supply

We observed significant changes in the elemental and intact polar lipid (IPL) composition of the archaeon Thermococcus kodakarensis (KOD1) in response to growth stage and phosphorus supply. Reducing the amount of organic supplements and phosphate in growth media resulted in significant decreases in cell size and cellular quotas of carbon (C), nitrogen (N), and phosphorus (P), which coincided with significant increases in cellular IPL quota and IPLs comprising multiple P atoms and hexose moieties. Relatively more cellular P was stored as IPLs in P-limited cells (2-8%) compared to control cells (<0.8%). We also identified a specific IPL biomarker containing a phosphatidyl-N-acetylhexoseamine headgroup that was relatively enriched during rapid cell division. These observations serve as empirical evidence of IPL adaptations in Archaea that will help to interpret the distribution of these biomarkers in natural systems. The reported cell quotas of C, N, and P represent the first such data for a specific archaeon and suggest that thermophiles are C-rich compared to the cell carbon-to-volume relationship reported for planktonic bacteria.

Effects of increasing seawater carbon dioxide concentrations on chain formation of the diatom Asterionellopsis glacialis

Diatoms can occur as single cells or as chain-forming aggregates. These two strategies affect buoyancy, predator evasion, light absorption and nutrient uptake. Adjacent cells in chains establish connections through various processes that determine strength and flexibility of the bonds, and at distinct cellular locations defining colony structure. Chain length has been found to vary with temperature and nutrient availability as well as being positively correlated with growth rate. However, the potential effect of enhanced carbon dioxide (CO2) concentrations and consequent changes in seawater carbonate chemistry on chain formation is virtually unknown. Here we report on experiments with semi-continuous cultures of the freshly isolated diatom Asterionellopsis glacialis grown under increasing CO2 levels ranging from 320 to 3400 µatm. We show that the number of cells comprising a chain, and therefore chain length, increases with rising CO2 concentrations. We also demonstrate that while cell division rate changes with CO2 concentrations, carbon, nitrogen and phosphorus cellular quotas vary proportionally, evident by unchanged organic matter ratios. Finally, beyond the optimum CO2 concentration for growth, carbon allocation changes from cellular storage to increased exudation of dissolved organic carbon. The observed structural adjustment in colony size could enable growth at high CO2 levels, since longer, spiral-shaped chains are likely to create microclimates with higher pH during the light period. Moreover increased chain length of Asterionellopsis glacialis may influence buoyancy and, consequently, affect competitive fitness as well as sinking rates. This would potentially impact the delicate balance between the microbial loop and export of organic matter, with consequences for atmospheric carbon dioxide.

CO2 and vitamin B12 interactions determine bioactive trace metal requirements of a subarctic Pacific diatom

Phytoplankton growth can be limited by numerous inorganic nutrients and organic growth factors. Using the subarctic diatom Attheya sp. in culture studies, we examined how the availability of vitamin B(12) and carbon dioxide partial pressure (pCO(2)) influences growth rate, primary productivity, cellular iron (Fe), cobalt (Co), zinc (Zn) and cadmium (Cd) quotas, and the net use efficiencies (NUEs) of these bioactive trace metals (mol C fixed per mol cellular trace metal per day). Under B(12)-replete conditions, cells grown at high pCO(2) had lower Fe, Zn and Cd quotas, and used those trace metals more efficiently in comparison with cells grown at low pCO(2). At high pCO(2), B(12)-limited cells had ~50% lower specific growth and carbon fixation rates, and used Fe ~15-fold less efficiently, and Zn and Cd ~3-fold less efficiently, in comparison with B(12)-replete cells. The observed higher Fe, Zn and Cd NUE under high pCO(2)/B(12)-replete conditions are consistent with predicted downregulation of carbon-concentrating mechanisms. Co quotas of B(12)-replete cells were 5- to 14-fold higher in comparison with B(12)-limited cells, suggesting that >80% of cellular Co of B(12)-limited cells was likely from B(12). Our results demonstrate that CO(2) and vitamin B(12) interactively influence growth, carbon fixation, trace metal requirements and trace metal NUE of this diatom. This suggests the need to consider complex feedback interactions between multiple environmental factors for this biogeochemically critical group of phytoplankton in the last glacial maximum as well as the current and future changing ocean.

Seawater carbonate chemistry and processes during experiments with Emiliania huxleyi (2005 Bergen) and Coccolithus braarudii (RCC 1200), 2010

The physiological performance of two coccolithophore species,Emiliania huxleyi and Coccolithus braarudii, was investigated during long-term exposure to elevated pCO2 levels. Mono-specific cultures were grown over 152 (E. huxleyi) and 65 (C. braarudii) generations while pCO2 was gradually increased to maximum levels of 1150 ?atm (E. huxleyi) and 930 ?atm (C. braarudii) and kept constant thereafter. Rates of cell growth and cell quotas of particulate organic carbon (POC), particulate inorganic carbon (PIC) and total particulate nitrogen (TPN) were determined repeatedly throughout the incubation period. Increasing pCO2 caused a decrease in cell growth rate of 9% and 29% in E. huxleyi and C. braarudii, respectively. In both species cellular PIC:TPN and PIC:POC ratios decreased in response to rising pCO2, whereas no change was observed in the POC:TPN ratios of E. huxleyi and C. braarudii. These results are consistent with those obtained in shorter-term high CO2exposure experiments following abrupt pertubations of the seawater carbonate system and indicate that for the strains tested here a gradual CO2 increase does not alleviate CO2/pH sensitivity.

Seawater carbonate chemistry and effects of CO2 and their modulation by light in the life-cycle stages of the coccolithophore Emiliania huxleyi strains RCC 1216 and 1217 during experiments, 2012

The effects of ocean acidification on the life-cycle stages of the coccolithophore Emiliania huxleyi and their by light were examined. Calcifying diploid and noncalcifying haploid cells (Roscoff culture collection 1216 and 1217) were acclimated to present-day and elevated CO2 partial pressures (PCO2; 38.5 vs. 101.3 Pa, ., 380 vs. 1000 matm) under low and high light (50 vs. 300 mmol photons m-2 s-1). Growth rates as well as quotas and production rates of C and N were measured. Sources of inorganic C for biomass buildup were using a 14C disequilibrium assay. Photosynthetic O2 evolution was measured as a function of dissolved inorganic C and light by means of membrane-inlet mass spectrometry. The diploid stage responded to elevated PCO2 by shunting resources from the production of particulate inorganic C toward organic C yet keeping the production of total particulate C constant. As the effect of ocean acidification was stronger under low light, the diploid stage might be less affected by increased acidity when energy availability is high. The haploid stage maintained elemental composition and production rates under elevated PCO2. Although both life-cycle stages involve different ways of dealing with elevated PCO2, the responses were generally modulated by energy availability, being typically most pronounced under low light. Additionally, PCO2 responses resembled those induced by high irradiances, indicating that ocean acidification affects the interplay between energy-generating processes (photosynthetic light reactions) and processes competing for energy (biomass buildup and calcification). A conceptual model is put forward explaining why the magnitude of single responses is determined by energy availability.

The effect of nitrate and phosphate availability on Emiliania huxleyi(NZEH) physiology under different CO2 scenarios

Growth and calcification of the marine coccolithophorid Emiliania huxleyi is affected by ocean acidification and macronutrients limitation and its response varies between strains. Here we investigated the physiological performance of a highly calcified E. huxleyi strain, NZEH, in a multiparametric experiment. Cells were exposed to different CO2 levels (ranging from 250 to 1314 µatm) under three nutrient conditions [nutrient replete (R), nitrate limited (-N), and phosphate limited (-P)]. We focused on calcite and organic carbon quotas and on nitrate and phosphate utilization by analyzing the activity of nitrate reductase (NRase) and alkaline phosphatase (APase), respectively. Particulate inorganic (PIC) and organic (POC) carbon quotas increased with increasing CO2 under R conditions but a different pattern was observed under nutrient limitation. The PIC:POC ratio decreased with increasing CO2 in nutrient limited cultures. Coccolith length increased with CO2 under all nutrient conditions but the coccosphere volume varied depending on the nutrient treatment. Maximum APase activity was found at 561 ?atm of CO2 (pH 7.92) in -P cultures and in R conditions, NRase activity increased linearly with CO2. These results suggest that E. huxleyi's competitive ability for nutrient uptake might be altered in future high-CO2 oceans. The combined dataset will be useful in model parameterizations of the carbon cycle and ocean acidification.

Seawater carbonate chemistry and biological processes of Emiliania huxleyi (PML B92/11) during experiments, 2011

The present study investigates the combined effect of phosphorous limitation, elevated partial pressure of CO2 (pCO2) and temperature on a calcifying strain of Emiliania huxleyi (PML B92/11) by means of a fully controlled continuous culture facility. Two levels of phosphorous limitation were consecutively applied by renewal of culture media (N:P = 26) at dilution rates (D) of 0.3 d- and 0.1 d-1. CO2 and temperature conditions were 300, 550 and 900 µatm pCO2 at 14 °C and 900 µatm pCO2 at 18 °C. In general, the steady state cell density and particulate organic carbon (POC) production increased with pCO2, yielding significantly higher concentrations in cultures grown at 900 µatm pCO2 compared to 300 and 550 µatm pCO2. At 900 µatm pCO2, elevation of temperature as expected for a greenhouse ocean, further increased cell densities and POC concentrations. In contrast to POC concentration, C-quotas (pmol C cell-1) were similar at D = 0.3 d-1 in all cultures. At D = 0.1 d-1, a reduction of C-quotas by up to 15% was observed in the 900 µatm pCO2 at 18 °C culture. As a result of growth rate reduction, POC:PON:POP ratios deviated strongly from the Redfield ratio, primarily due to an increase in POC. Ratios of particulate inorganic and organic carbon (PIC:POC) ranged from 0.14 to 0.18 at D = 0.3 d-1, and from 0.11 to 0.17 at D = 0.1 d-1, with variations primarily induced by the changes in POC. At D = 0.1 d-1, cell volume was reduced by up to 22% in cultures grown at 900 µatm pCO2. Our results indicate that changes in pCO2, temperature and phosphorus supply affect cell density, POC concentration and size of E. huxleyi (PML B92/11) to varying degrees, and will likely impact bloom development as well as biogeochemical cycling in a greenhouse ocean.