Experiments on polar seaweeds

Organisms populating benthic shallow water systems of both polar regions are adapted to a particularly harsh environment. We studied effects of freezing and the combination of high light intensities and low water temperatures on photosynthesis of key macroalgal species from the Arctic intertidal (Fucus distichus) and Antarctic subtidal (Palmaria decipiens). Photosynthetic activity of F. distichus specimens was monitored during the freezing process; there was a marked decrease in quantum yield with decreasing temperatures, and a rapid recovery as soon as temperatures increased again. Thus, under the experimental conditions tested, no indication of photodamage was found. Specimens of Palmaria were exposed to a combination of high light intensities and low water temperatures. A persistent impairment of photosynthetic activity occurred at 0°C at light intensities of 400 µmol photons m-2 s-1. In all treatments, there was a decreasing ratio of phycobiliproteins to chlorophyll a. Overall, the two studies provide baseline data for interpreting physiological responses of two important macroalgal species in an extreme environment, the polar coastal ecosystem.

Anemone abundance and productivity at North Vulcano Island in May 2011

Increased seawater pCO2, and in turn 'ocean acidification' (OA), is predicted to profoundly impact marine ecosystem diversity and function this century. Much research has already focussed on calcifying reef-forming corals (Class: Anthozoa) that appear particularly susceptible to OA via reduced net calcification. However, here we show that OA-like conditions can simultaneously enhance the ecological success of non-calcifying anthozoans, which not only play key ecological and biogeochemical roles in present day benthic ecosystems but also represent a model organism should calcifying anthozoans exist as less calcified (soft-bodied) forms in future oceans. Increased growth (abundance and size) of the sea anemone (Anemonia viridis) population was observed along a natural CO2 gradient at Vulcano, Italy. Both gross photosynthesis (PG) and respiration (R) increased with pCO2 indicating that the increased growth was, at least in part, fuelled by bottom up (CO2 stimulation) of metabolism. The increase of PG outweighed that of R and the genetic identity of the symbiotic microalgae (Symbiodinium spp.) remained unchanged (type A19) suggesting proximity to the vent site relieved CO2 limitation of the anemones' symbiotic microalgal population. Our observations of enhanced productivity with pCO2, which are consistent with previous reports for some calcifying corals, convey an increase in fitness that may enable non-calcifying anthozoans to thrive in future environments, i.e. higher seawater pCO2. Understanding how CO2-enhanced productivity of non- (and less-) calcifying anthozoans applies more widely to tropical ecosystems is a priority where such organisms can dominate benthic ecosystems, in particular following localized anthropogenic stress.

(Table 1) Productivity characteristics of phytoplankton and some concomitant factors in the Kara Sea in September 1993

Primary production in water column (P_p) varied from 107 to 312 mg C/m**2/day in Yenisey Bay: from 25 to 63 mg C/m**2/day in Obskaya Guba: and from 20 to 359 mg C/m**2/day in the open sea, that is: in the western Kara Sea and Ob-Yenisey shoals. The average concentration of chlorophyll a in the photosynthesis layer (C_ph) ranged from 0.2 to 1.8 mg/m**3 in these two regions, lower than in the estuaries of Ob (1.6-21.7 mg/m**3) and Yenisey (2.0-5.2 mg/m**3) Rivers. An inverse relation between surface salinity (S) and chlorophyll concentration (C_s) and chlorophyll concentration in the photosynthesis layer was found for all of the regions. The highest values of C_s and C_ph (0.8-22 mg/m**3) were measured at S<10 ppt, and the lowest values (0.2-0.8 mg/m**3) at S>22 ppt. A similar correlation of S with values of Pp occurred only in the Yenisey Bay and offshore regions. Obtained results agree well with the "outwelling" hypothesis. It states that large part of organic matter produced in estuaries is not used in estuarine trophic chains but is transported into adjacent sea areas and increases their productivity. Low values of Pp in the study regions may be attributed to such unfavorable factors as deficiency in nutrients, low temperature and turbidity, and lack of solar radiation.

(Table 1) Results of measurements of production characteristics of phytoplankton and some physical parameters of the environment in the northeastern Norwegian Sea in July 1995

From July 4 to 18,1995 surface chlorophyll a concentrations (C_cs) and integral primary production (C_ps) were studied in the northeastern part of the Norwegian Sea (73°42'N; 13°16'E), over a test area where an accident of the nuclear submarine Komsomolets had taken place. It was found that during this interval C_cs decreased by factor of about 3.3 (from 0.78 to 0.24 mg/m**3); average chlorophyll concentration within the photo-synthetic layer (C_cl) decreased by factor of about 3.5 (from 0.97 to 0.28 mg/m**3). The value of C_ps in the water column varied slightly (from 445 to 539 mg C/m**2 per day), since decrease in C_cl was compensated both by 1.5-fold growth of the photosynthetic layer thickness (from 40 to 60 m) and by 2.1-fold increase in its average assimilation number (from 0.58 to 1.20 mg C/mg chl a per hour). Monthly averages of C_ps were obtained from published data on seasonal C_cs changes and on the level of incident solar irradiation. They were found to be less than 100 mg C/m**2 per day in March and October and 100-500 mg C/m**2 per day in April-June. Annual primary production calculated from above values was equal to 105 g C/m**2.

Compositional and isotopic data on core MD84641

The organic carbon isotopic record of the sapropels(S1 and S3-S10) and intercalated marl oozes has been determined in a 12-m piston core from the eastern Mediterranean. The d13C_organic values are systematically lighter (mean=-21.0±0.82 per mil) in all sapropels and heavier (mean=-18.8±1.07 per mil) in the marl oozes. These differences are not due to variable marine and terrestrial organic matter mixtures because all values are heavier than modern plankton in the Mediterranean, there is no relationship between the C_organic/N ratios and the isotopic values, and published information on the abundance and distribution of organic biomarkers shows that terrestrial material constitutes a minor fraction of the total organic matter. Temperature effects on isotope fractionation are also discounted because the change in d13C_organic values between glacial and interglacial horizons is in the opposite sense. Diagenesis, which can produce relatively small changes in the carbon isotopic composition of sedimentary organic matter under certain circumstances, is unlikely to have caused the observed differences because this mechanism would cause an enrichmet in 12C, implying that all values were even heavier originally, and there is no secular trend in the d13C_organic record. The observed differences in d13C_organic between the two lithologies are probably produced by changes in the isotopic composition and the concentration of dissolved CO2. First, freshwater flooding during the formation of the sapropels caused the isotopic composition of the dissolved inorganic carbon in the surface waters of the Mediterranean to become lighter because of the 13C deficiency in fresh waters. Hence photosynthesis would have produced isotopically lighter organic material. Second, changes in atmospheric pCO2 between glacial and interglacial periods, as shown by the Vostok ice core, caused marked changes in the concentration of free dissolved CO2 in the mixed layer; lower values during glacial maxima caused a smaller fractionation of the carbon isotopes by phytoplankton, whereas levels were less limiting during the interglacials. Concentrations of dissolved CO2 could also have been much higher during the deposition of the sapropels because of the supply of regenerated CO2 to the mixed layer by upwelling, and this could have further lightened the d13C_organic values in the sapropels themselves. Carbon isotope records may provide an alternative method for estimating atmospheric pCO2 levels over longer time periods than can be obtained from ice cores.

Water chemistry of lakes in Zackenbergdalen between 1997-2003, East Greenland

The ecology of arctic lakes is strongly influenced by climate-generated variations in snow coverage and by the duration of the ice-free period, which, in turn, affect the physical and chemical conditions of the lakes (Wrona et al., 2005, http://www.acia.uaf.edu/PDFs/ACIA_Science_Chapters_Final/ACIA_Ch08_Final.pdf). Most arctic lakes are characterised by a long period (8-10 months) of ice-cover, cold water and low algal biomass. The water temperature and nutrient concentrations, and most probably the nutrient input from the catchments, are closely related to the duration of snow- and ice-cover in the lakes. In years when the ice-out is late, - that is, in late July, - phytoplankton photosynthesis is limited by the lack of light and nutrients. Less food is then available to the next link in the food chain, such as copepods and daphnids, with implication on their growth rates.

(Table 2) Production characteristics of phytoplankton and selected related factors at bottle sampling stations in the subtropical and tropical Atlantic Ocean

In October and November 2002, high and relatively high values of chlorophyll a concentration at the sea surface (Cchl) were observed in the English Channel (0.47 mg/m**3), in waters of the North Atlantic Current (0.25 mg/m**3 ), in the tropical and subtropical anticyclonic gyres (0.07-0.42 mg/m**3), and also in the southwestern region of the southern subtropical anticyclonic gyre (usually 0.11-0.23 mg/m**3). The central regions of the southern subtropical anticyclonic gyre (SATG) and the North Atlantic tropical gyre (NATR) were characterized by lower values of Cchl (0.02-0.08 mg/m**3 for the SATG and 0.07-0.14 mg/m**3 for the NATR). At most of the SATG stations, values of surface primary production (Cphs) varied from 2.5 to 5.5 mg C/m**3 per day and were mainly defined by fluctuations of Cchl (r = +0.78) rather than by those of the assimilation number (r = +0.54). Low assimilation activity of phytoplankton in these waters (1.3-4.6 mg chl a per hour) pointed to a lack of nutrients. Analysis of variability of their concentration and composition of photosynthetic pigments showed that, in waters north of 30°N, the growth of phytoplankton was mostly restricted by deficiency of nitrogen, while, in more southern areas, at the majority of stations (about 60%), phosphorus concentrations were minimal. At low concentrations of nitrates and nitrites, ammonium represented itself as a buffer that prevented planktonic algae from extreme degrees of nitric starvation. In tropical waters and in waters of the SATG, primary production throughout the water column varied from 240 to 380 mg C/m**2 30° per day. This level of productivity at stations with low values of C chl (<0.08 mg/m**3) was provided by a well-developed deep chlorophyll maximum and high transparency of water. Light curves of photosynthesis based on in situ measurements point to high efficiency of utilizing penetrating solar radiation by phytoplankton on cloudy days.

Morphological plasticity of the coral skeleton under CO2-driven seawater acidification

Ocean acidification causes corals to calcify at reduced rates, but current understanding of the underlying processes is limited. Here, we conduct a mechanistic study into how seawater acidification alters skeletal growth of the coral Stylophora pistillata. Reductions in colony calcification rates are manifested as increases in skeletal porosity at lower pH, while linear extension of skeletons remains unchanged. Inspection of the microstructure of skeletons and measurements of pH at the site of calcification indicate that dissolution is not responsible for changes in skeletal porosity. Instead, changes occur by enlargement of corallite-calyxes and thinning of associated skeletal elements, constituting a modification in skeleton architecture. We also detect increases in the organic matrix protein content of skeletons formed under lower pH. Overall, our study reveals that seawater acidification not only causes decreases in calcification, but can also cause morphological change of the coral skeleton to a more porous and potentially fragile phenotype.

Decreased light availability can amplify negative impacts of ocean acidification on calcifying coral reef organisms

Coral reef organisms are increasingly and simultaneously affected by global and local stressors such as ocean acidification (OA) and reduced light availability. However, knowledge of the interplay between OA and light availability is scarce. We exposed 2 calcifying coral reef species (the scleractinian coral Acropora millepora and the green alga Halimeda opuntia) to combinations of ambient and increased pCO2 (427 and 1073 µatm, respectively), and 2 light intensities (35 and 150 µmol photons/m**2/s) for 16 d. We evaluated the individual and combined effects of these 2 stressors on weight increase, calcification rates, O2 fluxes and chlorophyll a content for the species investigated. Weight increase of A. millepora was significantly reduced by OA (48%) and low light intensity (96%) compared to controls. While OA did not affect coral calcification in the light, it decreased calcification in the dark by 155%, leading to dissolution of the skeleton. H. opuntia weight increase was not affected by OA, but decreased (40%) at low light. OA did not affect algae calcification in the light, but decreased calcification in the dark by 164%, leading to dissolution. Low light significantly reduced gross photosynthesis (56 and 57%), net photosynthesis (62 and 60%) and respiration (43 and 48%) of A. millepora and H. opuntia, respectively. In contrast to A. millepora, H. opuntia significantly increased chlorophyll content by 15% over the course of the experiment. No interactive effects of OA and low light intensity were found on any response variable for either organism. However, A. millepora exhibited additive effects of OA and low light, while H. opuntia was only affected by low light. Thus, this study suggests that negative effects of low light and OA are additive on corals, which may have implications for management of river discharge into coastal coral reefs.

(Table 1) Results of measurements of productive characteristics of phytoplankton and environmental parameters in the Barents Sea in September-October 1997

The described studies were carried out in the eastern part of the sea during the end of the summer seasonal succession from September 1 to October 12, 1997. Concentration of chlorophyll a in the surface layer varied from 0.09 to 1.24 mg/m**3; it tended to increase in the southern regions (<74°N). Primary production in the water column (P_p) varied from 24 to 214 mg C/m**2/day and was on average 91 mg C/m**2/day. The low level of P_p seems to result from combination of physical and chemical environmental factors unfavorable for photosynthesis (e.g. deficiency of nutrients and low values of insolation and temperature) and intensive grazing of phytoplankton by zooplankton. The lower boundary of the photosynthetic layer in open waters was located at depth 60-75 m; irradiance there was 0.1-0.5% of incident irradiance. In deep-water regions (>200 m) the subsurface maximum of chlorophyll occurred in the layer at 20-40 m; usually this maximum resulted in formation of additional maxima of primary production.

Increased CO2 modifies the carbon balance and the photosynthetic yield of two common Arctic brown seaweeds: Desmarestia aculeata and Alaria esculenta

Ocean acidification affects with special intensity Arctic ecosystems, being marine photosynthetic organisms a primary target, although the consequences of this process in the carbon fluxes of Arctic algae are still unknown. The alteration of the cellular carbon balance due to physiological acclimation to an increased CO2 concentration (1300 ppm) in the common Arctic brown seaweeds Desmarestia aculeata and Alaria esculenta from Kongsfjorden (Svalbard) was analysed. Growth rate of D. aculeata was negatively affected by CO2 enrichment, while A. esculenta was positively affected, as a result of a different reorganization of the cellular carbon budget in both species. Desmarestia aculeata showed increased respiration, enhanced accumulation of storage biomolecules and elevated release of dissolved organic carbon, whereas A. esculenta showed decreased respiration and lower accumulation of storage biomolecules. Gross photosynthesis (measured both as O2 evolution and 14C fixation) was not affected in any of them, suggesting that photosynthesis was already saturated at normal CO2 conditions and did not participate in the acclimation response. However, electron transport rate changed in both species in opposite directions, indicating different energy requirements between treatments and species specificity. High CO2 levels also affected the N-metabolism, and 13C isotopic discrimination values from algal tissue pointed to a deactivation of carbon concentrating mechanisms. Since increased CO2 has the potential to modify physiological mechanisms in different ways in the species studied, it is expected that this may lead to changes in the Arctic seaweed community, which may propagate to the rest of the food web.