Seawater carbonate chemistry, growth rate and light sensitivity of marine diatom Chaetoceros muelleri (strain CSIRO CS-176) during experiments, 2011

This study investigated the impact of photon flux and elevated CO2 concentrations on growth and photosynthetic electron transport on the marine diatom Chaetoceros muelleri and looked for evidence for the presence of a CO2-concentrating mechanism (CCM). pH drift experiments clearly showed that C. muelleri has the capacity to use bicarbonate to acquire inorganic carbon through one or multiple CCMs. The final pH achieved in unbuffered cultures was not changed by light intensity, even under very low photon flux, implying a low energy demand of bicarbonate use via a CCM. In short-term pH drift experiments, only treatment with the carbonic anhydrase inhibitor ethoxyzolamide (EZ) slowed down the rise in pH considerably. EZ was also the only inhibitor that altered the final pH attained, although marginally. In growth experiments, CO2 availability was manipulated by changing the pH in closed flasks at a fixed dissolved inorganic carbon (DIC) concentration. Low-light-treated samples showed lower growth rates in elevated CO2conditions. No CO2 effect was recorded under high light exposure. The maximal photosynthetic capacity, however, increased with CO2 concentration in saturating, but not in subsaturating, light intensities. Growth and photosynthetic capacity therefore responded in opposite ways to increasing CO2 availability. The capacity to photoacclimate to high and low photon flux appeared not to be affected by CO2treatments. However, photoacclimation was restricted to growth photon fluxes between 30 and 300 µmol photons m-2 s-1. The light saturation points for photosynthetic electron transport and for growth coincided at 100 µmol photons m-2 s-1. Below 100 µmol photons m-2 s-1 the light saturation point for photosynthesis was higher than the growth photon flux (i.e. photosynthesis was not light saturated under growth conditions), whereas at higher growth photon flux, photosynthesis was saturated below growth light levels.

Seawater carbonate chemistry and growth, calcification of Thoracosphaera heimii in a laboratory experiment

Ocean acidification is considered a major threat to marine ecosystems and may particularly affect calcifying organisms such as corals, foraminifera and coccolithophores. Here we investigate the impact of elevated pCO2 and lowered pH on growth and calcification in the common calcareous dinoflagellate Thoracosphaera heimii. We observe a substantial reduction in growth rate, calcification and cyst stability of T. heimii under elevated pCO2. Furthermore, transcriptomic analyses reveal CO2 sensitive regulation of many genes, particularly those being associated to inorganic carbon acquisition and calcification. Stable carbon isotope fractionation for organic carbon production increased with increasing pCO2 whereas it decreased for calcification, which suggests interdependence between both processes. We also found a strong effect of pCO2 on the stable oxygen isotopic composition of calcite, in line with earlier observations concerning another T. heimii strain. The observed changes in stable oxygen and carbon isotope composition of T. heimii cysts may provide an ideal tool for reconstructing past seawater carbonate chemistry, and ultimately past pCO2. Although the function of calcification in T. heimii remains unresolved, this trait likely plays an important role in the ecological and evolutionary success of this species. Acting on calcification as well as growth, ocean acidification may therefore impose a great threat for T. heimii.

Experimental data on photophysiology and growth rates of Pararotalia calcariformata

The eastern Mediterranean is a hotspot of biological invasions. Numerous species of Indo-pacific origin have colonized the Mediterranean in recent times, including tropical symbiont-bearing foraminifera. Among these is the species Pararotalia calcariformata. Unlike other invasive foraminifera, this species has been discovered only two decades ago and is restricted to the eastern Mediterranean coast. Combining ecological, genetic and physiological observations, we attempt to explain the recent invasion of this species in the Mediterranean Sea. Using morphological and genetic data, we confirm the species attribution to P. calcariformata McCulloch 1977 and identify its symbionts as a consortium of diatom species dominated by Minutocellus polymorphus. We document photosynthetic activity of its endosymbionts using Pulse Amplitude Modulated Fluorometry and test the effects of elevated temperatures on growth rates of asexual offspring. The culturing of asexual offspring for 120 days shows a 30-day period of rapid growth followed by a period of slower growth. A subsequent 48-day temperature sensitivity experiment indicates a similar developmental pathway and high growth rate at 28°C, whereas an almost complete inhibition of growth was observed at 20°C and 35°C. This indicates that the offspring of this species may have lower tolerance to cold temperatures than what would be expected for species native to the Mediterranean. We expand this hypothesis by applying a Species Distribution Model (SDM) based on modern occurrences in the Mediterranean using three environmental variables: irradiance, turbidity and yearly minimum temperature. The model reproduces the observed restricted distribution and indicates that the range of the species will drastically expand westwards under future global change scenarios. We conclude that P. calcariformata established a population in the Levant because of the recent warming in the region. In line with observations from other groups of organisms, our results indicate that continued warming of the eastern Mediterranean will facilitate the invasion of more tropical marine taxa into the Mediterranean, disturbing local biodiversity and ecosystem structure.

Relative sensitivity of soft-bottom intertidal macrofauna to increased CO2 and experimental stress

We used a controlled CO2 perturbation experiment to test hypotheses about changes in diversity, composition and structure of soft-bottom intertidal macrobenthic assemblages, under realistic and locally relevant scenarios of seawater acidification. Patches of undisturbed sediment were collected from 2 types of intertidal sedimentary habitat in the Ria Formosa coastal lagoon (South Portugal) and exposed to 2 levels of seawater acidification (pH reduced by 0.3 and 0.6 units) and 1 unmanipulated (control) level. After 75 d the assemblages differed significantly between the 2 types of sediment and between field controls and the ex situ treatments, but not among the 3 pH levels tested. The naturally high values of total alkalinity buffered seawater from the changes imposed on carbonate chemistry and may have contributed to offsetting acidification at the local scale. Observed differences on biota were strongly related to the organic matter content and grain-size of the sediments, particularly to the fractions of medium and coarse sand. Soft-bottom intertidal macrofauna was significantly affected by the stress of being held in an artificial environment, but not by CO2-induced seawater acidification. Given the previously observed variations in the sensitivities of marine organisms to seawater acidification, direct extrapolations of the present findings to different regions or other types of assemblages do not seem advisable. However, the contribution of ex situ studies to the assessment of ecosystem-level responses to environmental disturbances could generally be improved by incorporating adequate field controls in the experimental design.

Temperature modulates coccolithophorid sensitivity of growth, photosynthesis and calcification to increasing seawater pCO2

Increasing atmospheric CO2 concentrations are expected to impact pelagic ecosystem functioning in the near future by driving ocean warming and acidification. While numerous studies have investigated impacts of rising temperature and seawater acidification on planktonic organisms separately, little is presently known on their combined effects. To test for possible synergistic effects we exposed two coccolithophore species, Emiliania huxleyi and Gephyrocapsa oceanica, to a CO2 gradient ranging from ~0.5-250 µmol/kg (i.e. ~20-6000 µatm pCO2) at three different temperatures (i.e. 10, 15, 20°C for E. huxleyi and 15, 20, 25°C for G. oceanica). Both species showed CO2-dependent optimum-curve responses for growth, photosynthesis and calcification rates at all temperatures. Increased temperature generally enhanced growth and production rates and modified sensitivities of metabolic processes to increasing CO2. CO2 optimum concentrations for growth, calcification, and organic carbon fixation rates were only marginally influenced from low to intermediate temperatures. However, there was a clear optimum shift towards higher CO2 concentrations from intermediate to high temperatures in both species. Our results demonstrate that the CO2 concentration where optimum growth, calcification and carbon fixation rates occur is modulated by temperature. Thus, the response of a coccolithophore strain to ocean acidification at a given temperature can be negative, neutral or positive depending on that strain's temperature optimum. This emphasizes that the cellular responses of coccolithophores to ocean acidification can only be judged accurately when interpreted in the proper eco-physiological context of a given strain or species. Addressing the synergistic effects of changing carbonate chemistry and temperature is an essential step when assessing the success of coccolithophores in the future ocean.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.