Kim I. Currie

TESTING THE EFFECTS OF OCEAN ACIDIFICATION ON ALGAL METABOLISM: CONSIDERATIONS FOR EXPERIMENTAL DESIGNS(1).

Ocean acidification describes changes in the carbonate chemistry of the ocean due to the increased absorption of anthropogenically released CO2. Experiments to elucidate the biological effects of ocean acidification on algae are not straightforward because when pH is altered, the carbon speciation in seawater is altered, which has implications for photosynthesis and, for calcifying algae, calcification. Furthermore, photosynthesis, respiration, and calcification will themselves alter the pH of the seawater medium. In this review, algal physiologists and seawater carbonate chemists combine their knowledge to provide the fundamental information on carbon physiology and seawater carbonate chemistry required to comprehend the complexities of how ocean acidification might affect algae metabolism. A wide range in responses of algae to ocean acidification has been observed, which may be explained by differences in algal physiology, timescales of the responses measured, study duration, and the method employed to alter pH. Two methods have been widely used in a range of experimental systems: CO2 bubbling and HCl/NaOH additions. These methods affect the speciation of carbonate ions in the culture medium differently; we discuss how this could influence the biological responses of algae and suggest a third method based on HCl/NaHCO3 additions. We then discuss eight key points that should be considered prior to setting up experiments, including which method of manipulating pH to choose, monitoring during experiments, techniques for adding acidified seawater, biological side effects, and other environmental factors. Finally, we consider incubation timescales and prior conditioning of algae in terms of regulation, acclimation, and adaptation to ocean acidification.

Ocean acidification at high latitudes: potential effects on functioning of the Antarctic bivalve Laternula elliptica.

Ocean acidification is a well recognised threat to marine ecosystems. High latitude regions are predicted to be particularly affected due to cold waters and naturally low carbonate saturation levels. This is of concern for organisms utilising calcium carbonate (CaCO3) to generate shells or skeletons. Studies of potential effects of future levels of pCO2 on high latitude calcifiers are at present limited, and there is little understanding of their potential to acclimate to these changes. We describe a laboratory experiment to compare physiological and metabolic responses of a key benthic bivalve, Laternula elliptica, at pCO2 levels of their natural environment (430 µatm, pH 7.99; based on field measurements) with those predicted for 2100 (735 µatm, pH 7.78) and glacial levels (187 µatm, pH 8.32). Adult L. elliptica basal metabolism (oxygen consumption rates) and heat shock protein HSP70 gene expression levels increased in response both to lowering and elevation of pH. Expression of chitin synthase (CHS), a key enzyme involved in synthesis of bivalve shells, was significantly up-regulated in individuals at pH 7.78, indicating L. elliptica were working harder to calcify in seawater undersaturated in aragonite (ΩAr = 0.71), the CaCO3 polymorph of which their shells are comprised. The different response variables were influenced by pH in differing ways, highlighting the importance of assessing a variety of factors to determine the likely impact of pH change. In combination, the results indicate a negative effect of ocean acidification on whole-organism functioning of L. elliptica over relatively short terms (weeks-months) that may be energetically difficult to maintain over longer time periods. Importantly, however, the observed changes in L. elliptica CHS gene expression provides evidence for biological control over the shell formation process, which may enable some degree of adaptation or acclimation to future ocean acidification scenarios.

CARBON-USE STRATEGIES IN MACROALGAE: DIFFERENTIAL RESPONSES TO LOWERED PH AND IMPLICATIONS FOR OCEAN ACIDIFICATION1

Ocean acidification (OA) is a reduction in oceanic pH due to increased absorption of anthropogenically produced CO2. This change alters the seawater concentrations of inorganic carbon species that are utilized by macroalgae for photosynthesis and calcification: CO2 and HCO3− increase; CO32− decreases. Two common methods of experimentally reducing seawater pH differentially alter other aspects of carbonate chemistry: the addition of CO2 gas mimics changes predicted due to OA, while the addition of HCl results in a comparatively lower [HCO3−]. We measured the short‐term photosynthetic responses of five macroalgal species with various carbon‐use strategies in one of three seawater pH treatments: pH 7.5 lowered by bubbling CO2 gas, pH 7.5 lowered by HCl, and ambient pH 7.9. There was no difference in photosynthetic rates between the CO2, HCl, or pH 7.9 treatments for any of the species examined. However, the ability of macroalgae to raise the pH of the surrounding seawater through carbon uptake was greatest in the pH 7.5 treatments. Modeling of pH change due to carbon assimilation indicated that macroalgal species that could utilize HCO3− increased their use of CO2 in the pH 7.5 treatments compared to pH 7.9 treatments. Species only capable of using CO2 did so exclusively in all treatments. Although CO2 is not likely to be limiting for photosynthesis for the macroalgal species examined, the diffusive uptake of CO2 is less energetically expensive than active HCO3− uptake, and so HCO3−‐using macroalgae may benefit in future seawater with elevated CO2.

Diurnal fluctuations in seawater pH influence the response of a calcifying macroalga to ocean acidification

Coastal ecosystems that are characterized by kelp forests encounter daily pH fluctuations, driven by photosynthesis and respiration, which are larger than pH changes owing to ocean acidification (OA) projected for surface ocean waters by 2100. We investigated whether mimicry of biologically mediated diurnal shifts in pH—based for the first time on pH time-series measurements within a kelp forest—would offset or amplify the negative effects of OA on calcifiers. In a 40-day laboratory experiment, the calcifying coralline macroalga, Arthrocardia corymbosa, was exposed to two mean pH treatments (8.05 or 7.65). For each mean, two experimental pH manipulations were applied. In one treatment, pH was held constant. In the second treatment, pH was manipulated around the mean (as a step-function), 0.4 pH units higher during daylight and 0.4 units lower during darkness to approximate diurnal fluctuations in a kelp forest. In all cases, growth rates were lower at a reduced mean pH, and fluctuations in pH acted additively to further reduce growth. Photosynthesis, recruitment and elemental composition did not change with pH, but δ13C increased at lower mean pH. Including environmental heterogeneity in experimental design will assist with a more accurate assessment of the responses of calcifiers to OA.

Apparatus for continuous-flow underway spectrophotometric measurement of surface water pH

A continuous-flow method for spectrophotometric measurement of seawater pH providing high temporal resolution has been developed incorporating a CCD-based spectrophotometer and dual pumps for seawater and indicator dye solution. The method showed good laboratory precision (±0.0007 pH units). The accuracy of the measurement system was assessed by comparing in situ measured pCO 2 values calculated from measured pH and alkalinity. An average difference between the results of ca. ± 8 μatm was obtained. Some difficulties was encountered during longer-term deployment on offshore cruises, including particulate matter in the surface seawater line, bio-fouling of the optical cell, and air-bubbles in the surface seawater line. Protocols were adopted to either prevent these problems or to recognise and remove data affected by them. Accuracy was also assessed by comparing the measured pH with values obtained by conventional discrete spectrophotometric measurements. On the first cruise, poor accuracy was found during potion of the cruise where pH < 7.80 was recorded by the continuous-flow apparatus, probably because of fouling of the pump and connector tubing, which was very evident on this cruise. In other regions, the agreement between the two methods was within ± 0.02 pH units. On the second offshore cruise, accuracy was assessed by analysing both a continuous flow of surface seawater and an isolated seawater sample that had been atmospherically equilibrated. In the first case, an accuracy of ± 0.005 pH units was achieved, while the second case gave a better result of ± 0.002 pH units. We conclude that after minor improvements, the measurement system should be well suited to high-frequency pH measurements of surface waters.

Oceanic fronts: transition zones for bacterioplankton community composition

Oceanic fronts are widespread mesoscale features that exist in the boundary between different water masses. Despite the recognized importance of bacterioplankton (including bacteria and archaea) on the marine biogeochemical cycles and the ubiquitousness of fronts, the effect of frontal zones on the distribution of bacterioplankton community remains unknown. Using 16S rRNA gene sequencing coupled with a high spatial resolution analysis of the physical properties of the water masses, we demonstrate strong shifts in bacterioplankton community composition (BCC) across the subtropical frontal zone off New Zealand. The transition between water masses resulted in a clear modification of the dominant taxa and a significant increase in community dissimilarity. Our results, linking physical oceanography and marine molecular ecology, support the strong role of oceanic frontal zones in delimiting the distribution of bacterioplankton in the ocean.

Ocean acidification in New Zealand waters: trends and impacts

ABSTRACT The threat posed by ocean acidification (OA) to the diversity and productivity of New Zealand marine ecosystems is assessed in a synthesis of published trends and impacts. A 20-year time series in Subantarctic water, and a national coastal monitoring programme, provide insight into pH variability, and context for experimental design, modelling and projections. A review of the potential impact of changes in the carbonate system on the major phyla in New Zealand waters confirms international observations that calcifying organisms, and particularly their early life-history stages, are vulnerable. The synthesis considers ecosystem and socio-economic impacts, and identifies current knowledge gaps and future research directions, including mechanistic studies of OA sensitivity. Advanced ecosystem models of OA, that incorporate the indirect effects of OA and interactions with other climate stressors, are required for robust projection of the future status of New Zealand marine ecosystems.

Climate change projections for the surface ocean around New Zealand

ABSTRACT The future status of the surface ocean around New Zealand was projected using two Earth System Models and four emission scenarios. By 2100 mean changes are largest under Representative Concentration Pathway 8.5 (RCP8.5), with a +2.5°C increase in sea surface temperature, and decreases in surface mixed layer depth (15%), macronutrients (7.5–20%), primary production (4.5%) and particle flux (12%). Largest macronutrient declines occur in the eastern Chatham Rise and subantarctic waters to the south, whereas dissolved iron increases in subtropical waters. Surface pH projections, validated against subantarctic time-series data, indicate a 0.335 decline to ∼7.77 by 2100. However, projected pH is sensitive to future CO2 emissions, remaining within the current range under RCP2.6, but decreasing below it by 2040 with all other scenarios. Sub-regions vulnerable to climate change include the Chatham Rise, polar waters south of 50°S, and subtropical waters north of New Zealand, whereas the central Tasman Sea is least affected.

Late‐summer biogeochemistry in the Mertz Polynya: East Antarctica

A marked reconfiguration of the Mertz Polynya following the 2010 calving of the Mertz Glacier Tongue has been associated with a decrease in the size and activity of the polynya. We report observations of the oceanic carbonate (CO 2 ) system in late-summer 2013, the third post-calving summer season. Estimates of seasonal net community production (NCP) based on inorganic carbon deficits and the oxygen-argon ratio indicate that the waters on the shelf to the east of Commonwealth Bay (adjacent to the Mertz Glacier) remain productive compared to pre-calving conditions. The input of residual or excess alkalinity from melting sea ice is found to contribute to the seasonal enhancement of carbonate saturation state and pH in shelf waters. Mean rates of NCP in 2012–2013 are more than twice as large as those observed in the pre-calving summers of 2001 and 2008 and suggest that the new (post-calving) configuration of the polynya favors enhanced net community production and a stronger surface ocean sink for atmospheric CO 2 due at least in part to the redistribution of sea ice and associated changes in summer surface stratification.

Microalgal photophysiology and macronutrient distribution in summer sea ice in the Amundsen and Ross Seas, Antarctica

Our study addresses how environmental variables, such as macronutrients concentrations, snow cover, carbonate chemistry and salinity affect the photophysiology and biomass of Antarctic sea-ice algae. We have measured vertical profiles of inorganic macronutrients (phosphate, nitrite + nitrate and silicic acid) in summer sea ice and photophysiology of ice algal assemblages in the poorly studied Amundsen and Ross Seas sectors of the Southern Ocean. Brine-scaled bacterial abundance, chl a and macronutrient concentrations were often high in the ice and positively correlated with each other. Analysis of photosystem II rapid light curves showed that microalgal cells in samples with high phosphate and nitrite + nitrate concentrations had reduced maximum relative electron transport rate and photosynthetic efficiency. We also observed strong couplings of PSII parameters to snow depth, ice thickness and brine salinity, which highlights a wide range of photoacclimation in Antarctic pack-ice algae. It is likely that the pack ice was in a post-bloom situation during the late sea-ice season, with low photosynthetic efficiency and a high degree of nutrient accumulation occurring in the ice. In order to predict how key biogeochemical processes are affected by future changes in sea ice cover, such as in situ photosynthesis and nutrient cycling, we need to understand how physicochemical properties of sea ice affect the microbial community. Our results support existing hypothesis about sea-ice algal photophysiology, and provide additional observations on high nutrient concentrations in sea ice that could influence the planktonic communities as the ice is retreating.