Near-bottom zooplankton aggregations in Kongsfjorden with link to images from the optical zooplankton sensor MOKI

Near-bottom zooplankton communities have rarely been studied despite numerous reports of high zooplankton concentrations, probably due to methodological constraints. In Kongsfjorden, Svalbard, the near-bottom layer was studied for the first time by combining daytime deployments of a remotely operated vehicle (ROV), the optical zooplankton sensor moored on-sight key species investigation (MOKI), and Tucker trawl sampling. ROV data from the fjord entrance and the inner fjord showed high near-bottom abundances of euphausiids with a mean concentration of 17.3 ± 3.5 n/100 m**3. With the MOKI system, we observed varying numbers of euphausiids, amphipods, chaetognaths, and copepods on the seafloor at six stations. Light-induced zooplankton swarms reached densities in the order of 90,000 (euphausiids), 120,000 (amphipods), and 470,000 ind/m**3 (chaetognaths), whereas older copepodids of Calanus hyperboreus and C. glacialis did not respond to light. They were abundant at the seafloor and 5 m above and showed maximum abundance of 65,000 ind/m**3. Tucker trawl data provided an overview of the seasonal vertical distribution of euphausiids. The most abundant species Thysanoessa inermis reached near-bottom concentrations of 270 ind/m**3. Regional distribution was neither related to depth nor to location in the fjord. The taxa observed were all part of the pelagic community. Our observations suggest the presence of near-bottom macrozooplankton also in other regions and challenge the current view of bentho-pelagic coupling. Neglecting this community may cause severe underestimates of the stock of elagic zooplankton, especially predatory species, which link secondary production with higher trophic levels.

Oxygen, pH and calcium dynamics and fluxes at the thallus surface under ambient (8.1) and low (7.8) seawater pH and across a range of irradiances

Presently, an incomplete mechanistic understanding of tropical reef macroalgae photosynthesis and calcification restricts predictions of how these important autotrophs will respond to global change. Therefore, we investigated the mechanistic link between inorganic carbon uptake pathways, photosynthesis and calcification in a tropical crustose coralline alga (CCA) using microsensors. We measured pH, oxygen (O2), and calcium (Ca2+) dynamics and fluxes at the thallus surface under ambient (8.1) and low (7.8) seawater pH (pHSW) and across a range of irradiances. Acetazolamide (AZ) was used to inhibit extracellular carbonic anhydrase (CAext), which mediates hydrolysis of HCO3-, and 4,4' diisothiocyanatostilbene-2,2'-disulphonate (DIDS) that blocks direct HCO3- uptake by anion exchange transport. Both inhibited photosynthesis, suggesting both diffusive uptake of CO2 via HCO3- hydrolysis to CO2 and direct HCO3- ion transport are important in this CCA. Surface pH was raised approximately 0.3 units at saturating irradiance, but less when CAext was inhibited. Surface pH was lower at pHSW 7.8 than pHSW 8.1 in the dark, but not in the light. The Ca2+ fluxes were large, complex and temporally variable, but revealed net Ca2+ uptake under all conditions. The temporal variability in Ca2+ dynamics was potentially related to localized dissolution during epithallial cell sloughing, a strategy of CCA to remove epiphytes. Simultaneous Ca2+ and pH dynamics suggest the presence of Ca2+/H+ exchange. Rapid light-induced H+ surface dynamics that continued after inhibition of photosynthesis revealed the presence of a light-mediated, but photosynthesis-independent, proton pump. Thus, the study indicates metabolic control of surface pH can occur in CCA through photosynthesis and light-inducible H+ pumps. Our results suggest that complex light-induced ion pumps play an important role in biological processes related to inorganic carbon uptake and calcification in CCA.

Seawater carbonate chemistry, chlorophyll a and photosynthetic carbon fixation rate under different light and pCO2 conditions during experiments, 2010

We carried out short term pCO2/pH perturbation experiments in the coastal waters of the South China Sea to evaluate the combined effects of seawater acidification (low pH/high pCO2) and solar UV radiation (UVR, 280-400 nm) on photosynthetic carbon fixation of phytoplankton assemblages. Under photosynthetically active radiation (PAR) alone treatments, reduced pCO2 (190 ppmv) with increased pH resulted in a significant decrease in the photosynthetic carbon fixation rate (about 23%), while enriched pCO2 (700 ppmv) with lowered pH had no significant effect on the photosynthetic performance compared to the ambient level. The apparent photosynthetic efficiency decreased under the reduced pCO2 level, probably due to C-limitation as well as energy being diverged for up-regulation of carbon concentrating mechanisms (CCMs). In the presence of UVR, both UV-A and UV-B caused photosynthetic inhibition, though UV-A appeared to enhance the photosynthetic efficiency under lower PAR levels. UV-B caused less inhibition of photosynthesis under the reduced pCO2 level, probably because of its contribution to the inorganic carbon (Ci)-acquisition processes. Under the seawater acidification conditions (enriched pCO2), both UV-A and UV-B reduced the photosynthetic carbon fixation to higher extents compared to the ambient pCO2 conditions. We conclude that solar UV and seawater acidification could synergistically inhibit photosynthesis.

The photo-physiological response of a model cnidarian-dinoflagellate symbiosis to CO2-induced acidification at the cellular level

We measured the relationship between CO2-induced seawater acidification, photo-physiological performance and intracellular pH (pHi) in a model cnidarian-dinoflagellate symbiosis - the sea anemone Aiptasia sp. -under ambient (289.94 ± 12.54 µatm), intermediate (687.40 ± 25.10 µatm) and high (1459.92 ± 65.51 µatm) CO2 conditions. These treatments represented current CO2 levels, in addition to CO2 stabilisation scenarios IV and VI provided by the Intergovernmental Panel on Climate Change (IPCC). Anemones were exposed to each treatment for two months and sampled at regular intervals. At each time-point we measured a series of physiological responses: maximum dark-adapted fluorescent yield of PSII (Fv/Fm), gross photosynthetic rate, respiration rate, symbiont population density, and light-adapted pHi of both the dinoflagellate symbiont and isolated host anemone cell. We observed increases in all but one photo-physiological parameter (Pgross:R ratio). At the cellular level, increases in light-adapted symbiont pHi were observed under both intermediate and high CO2 treatments, relative to control conditions (pHi 7.35 and 7.46 versus pHi 7.25, respectively). The response of light-adapted host pHi was more complex, however, with no change observed under the intermediate CO2 treatment, but a 0.3 pH-unit increase under the high CO2 treatment (pHi 7.19 and 7.48, respectively). This difference is likely a result of a disproportionate increase in photosynthesis relative to respiration at the higher CO2 concentration. Our results suggest that, rather than causing cellular acidosis, the addition of CO2 will enhance photosynthetic performance, enabling both the symbiont and host cell to withstand predicted ocean acidification scenarios.

Light availability determines susceptibility of reef building corals to ocean acidification

Elevated seawater pCO2, and in turn ocean acidification (OA), is now widely acknowledged to reduce calcification and growth of reef building corals. As with other environmental factors (e.g., temperature and nutrients), light availability fundamentally regulates calcification and is predicted to change for future reef environments alongside elevated pCO2 via altered physical processes (e.g., sea level rise and turbidity); however, any potential role of light in regulating the OA-induced reduction of calcification is still unknown. We employed a multifactorial growth experiment to determine how light intensity and pCO2 together modify calcification for model coral species from two key genera, Acropora horrida and Porites cylindrica, occupying similar ecological niches but with different physiologies. We show that elevated pCO2 (OA)-induced losses of calcification in the light (G L) but not darkness (G D) were greatest under low-light growth conditions, in particular for A. horrida. High-light growth conditions therefore dampened the impact of OA upon G L but not G D. Gross photosynthesis (P G) responded in a reciprocal manner to G L suggesting OA-relieved pCO2 limitation of P G under high-light growth conditions to effectively enhance G L. A multivariate analysis of past OA experiments was used to evaluate whether our test species responses were more widely applicable across their respective genera. Indeed, the light intensity for growth was identified as a significant factor influencing the OA-induced decline of calcification for species of Acropora but not Porites. Whereas low-light conditions can provide a refuge for hard corals from thermal and light stress, our study suggests that lower light availability will potentially increase the susceptibility of key coral species to OA.

CO2-induced ocean acidification increases anxiety in Rockfish via alteration of GABAA receptor functioning

The average surface pH of the ocean is dropping at a rapid rate due to the dissolution of anthropogenic CO2, raising concerns for marine life. Additionally, some coastal areas periodically experience upwelling of CO2-enriched water with reduced pH. Previous research has demonstrated ocean acidification (OA)-induced changes in behavioural and sensory systems including olfaction, which is due to altered function of neural gamma-aminobutyric acid type A (GABAA) receptors. Here, we used a camera-based tracking software system to examine whether OA-dependent changes in GABAA receptors affect anxiety in juvenile Californian rockfish (Sebastes diploproa). Anxiety was estimated using behavioural tests that measure light/dark preference (scototaxis) and proximity to an object. After one week in OA conditions projected for the next century in the California shore (1125 ± 100 µatm, pH 7.75), anxiety was significantly increased relative to controls (483 ± 40 µatm CO2, pH 8.1). The GABAA-receptor agonist muscimol, but not the antagonist gabazine, caused a significant increase in anxiety consistent with altered Cl- flux in OA-exposed fish. OA-exposed fish remained more anxious even after 7 days back in control seawater; however, they resumed their normal behaviour by day 12. These results show that OA could severely alter rockfish behaviour; however, this effect is reversible.

Alleviation of solar ultraviolet radiation (UVR)-induced photoinhibition in diatom Chaetoceros curvisetus by ocean acidification

The study aimed to unravel the interaction between ocean acidification and solar ultraviolet radiation (UVR) in Chaetoceros curvisetus. Chaetoceros curvisetus cells were acclimated to high CO2 (HC, 1000 ppmv) and low CO2 concentration (control, LC, 380 ppmv) for 14 days. Cell density, specific growth rate and chlorophyll were measured. The acclimated cells were then exposed to PAB (photosynthetically active radiation (PAR) + UV-A + UV-B), PA (PAR + UV-A) or P (PAR) for 60 min. Photochemical efficiency (phi PSII), relative electron transport rate (rETR) and the recovery of ?PSII were determined. HC induced higher cell density and specific growth rate compared with LC. However, no difference was found in chlorophyll between HC and LC. Moreover, phi PSII and rETRs were higher under HC than LC in response to solar UVR. P exposure led to faster recovery of phi PSII, both under HC and LC, than PA and PAB exposure. It appeared that harmful effects of UVR on C. curvisetus could be counteracted by ocean acidification simulated by high CO2 when the effect of climate change is not beyond the tolerance of cells.

Abundance and biomass of protozooplankton from the iron-induced phytoplankton bloom during the EisenEx experiment in 2000 in the Southern Ocean

The responses of larger (>50 µm in diameter) protozooplankton groups to a phytoplankton bloom induced by in situ iron fertilization (EisenEx) in the Polar Frontal Zone (PFZ) of the Southern Ocean in austral spring are presented. During the 21 days of the experiment, samples were collected from seven discrete depths in the upper 150 m inside and outside the fertilized patch for the enumeration of acantharia, foraminifera, radiolaria, heliozoa, tintinnid ciliates and aplastidic thecate dinoflagellates. Inside the patch, acantharian numbers increased twofold, but only negligibly in surrounding waters. This finding is of major interest, since acantharia are suggested to be involved in the formation of barite (BaSO_4 ) found in sediments and which is a palaeoindicator of both ancient and modern high productivity regimes. Foraminifera increased significantly in abundance inside and outside the fertilized patch. However the marked increase of juveniles after a full moon event suggests a lunar periodicity in the reproduction cycle of some foraminiferan species rather than a reproductive response to enhanced food availability. In contrast, adult radiolaria showed no clear trend during the experiment, but juveniles increased threefold indicating elevated reproduction. Aplastidic thecate dinoflagellates almost doubled in numbers and biomass, but also increased outside the patch. Tintinnid numbers decreased twofold, although biomass remained constant due to a shift in the size spectrum. Empty tintinnid loricae, however, increased by a factor of two indicating that grazing pressure on this group mainly by copepods intensified during EisenEx. The results show that iron-fertilization experiments can shed light on the biology and the role of these larger protists in pelagic ecosystem which will improve their use as proxies in palaeoceanography.

Combined effects of solar UV radiation and CO2-induced seawater acidification on photosynthetic carbon fixation of phytoplankton assemblages in the South China Sea, 2010

We carried out short term pCO2/pH perturbation experiments in the coastal waters of the South China Sea to evaluate the combined effects of seawater acidification (low pH/high pCO2) and solar UV radiation (UVR, 280-400 nm) on photosynthetic carbon fixation of phytoplankton assemblages. Under photosynthetically active radiation (PAR) alone treatments, reduced pCO2 (190 ppmv) with increased pH resulted in a significant decrease in the photosynthetic carbon fixation rate (about 23%), while enriched pCO2 (700 ppmv) with lowered pH had no significant effect on the photosynthetic performance compared to the ambient level. The apparent photosynthetic efficiency decreased under the reduced pCO2 level, probably due to C-limitation as well as energy being diverged for up-regulation of carbon concentrating mechanisms (CCMs). In the presence of UVR, both UV-A and UV-B caused photosynthetic inhibition, though UV-A appeared to enhance the photosynthetic efficiency under lower PAR levels. UV-B caused less inhibition of photosynthesis under the reduced pCO2 level, probably because of its contribution to the inorganic carbon (Ci)-acquisition processes. Under the seawater acidification conditions (enriched pCO2), both UV-A and UV-B reduced the photosynthetic carbon fixation to higher extents compared to the ambient pCO2 conditions. We conclude that solar UV and seawater acidification could synergistically inhibit photosynthesis.

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.