Photosynthetic activity buffers ocean acidification in seagrass meadows

Macrophytes growing in shallow coastal zones characterised by intense metabolic activity have the capacity to modify pH within their canopy and beyond. We observed diel pH changes in shallow (5-12 m) seagrass (Posidonia oceanica) meadows spanning 0.06 pH units in September to 0.24 units in June. The carbonate system (pH, DIC, and aragonite saturation state (omega Ar)) and O2 within the meadows displayed strong diel variability driven by primary productivity, and changes in chemistry were related to structural parameters of the meadow, in particular, the leaf surface area available for photosynthesis (LAI). LAI was positively correlated to mean, max and range pHNBS and max and range omega Ar. In June, vertical mixing (as Turbulent Kinetic Energy) influenced max and min omega Ar, while in September there was no effect of hydrodynamics on the carbonate system within the canopy. Max and range omega Ar within the meadow showed a positive trend with the calcium carbonate load of the leaves, pointing to a possible link between structural parameters, omega Ar and carbonate deposition.

Physicochemical sea ice and water characteristics near Point Barrow 2005-2006

Photosynthetic parameters of phytoplankton and sea ice algae from landfast sea ice of the Chukchi Sea off Point Barrow, Alaska, were assessed in spring 2005 and winter through spring 2006 using Pulse Amplitude Modulated (PAM) fluorometry including estimates of maximum quantum efficiency (Fv/Fm), maximum relative electron transport rate (rETRmax), photosynthetic efficiency (alpha), and the photoadaptive index (Ek). The use of centrifuged brine samples allowed to document vertical gradients in ice algal acclimation with 5 cm vertical resolution for the first time. Bottom ice algae (0-5 cm from ice-water interface) expressed low Fv/Fm (0.331-0.426) and low alpha (0.098-0.130 /(µmol photons/m**2/s)) in December. Fv/Fm and alpha increased in March and May (0.468-0.588 and 0.141-0.438 /(µmol photons/m**2/s), respectively) indicating increased photosynthetic activity. In addition, increases in rETRmax (3.3-16.4 a.u.) and Ek (20-88 µmol photons/m**2/s) from December to May illustrates a higher potential for primary productivity as communities become better acclimated to under-ice light conditions. In conclusion, photosynthetic performance by ice algae (as assessed by PAM fluorometry) was tightly linked to sea ice salinity, temperature, and inorganic nutrient concentrations (mainly nitrogen).

Phytoplankton in the Nhatrang Bay, South China Sea

Species composition, phytoplankton abundance, and relative yield of variable fluorescence (F_v/F_m) were determined in the mesotrophic Nhatrang Bay in October-November 2004. Species diversity (250 taxonomic units) and heterogeneity of the phytoplankton structure were high. With respect to number of species and their abundance, diatoms prevailed. In selected parts of the bay, dinoflagellates dominated. Average biomass in the water column under 1 m**2 (Bt) varied from 2.3 to 64.4 mg C/m**3 (av. 31.0 mg C/m**3). Bt values were the lowest at stations nearest to the river mouth. Seaward, Bt increased. Bt values increased with depth at some stations and decreased at others. In surface layers biomass was lower than that in the underlying waters. F_v/F_m values ranged from 0.10 to 0.64 (av. 0.49). The lowest F_v/F_m values were observed in the area close to the seaport. Over greater part of the bay, F_v/F_m values were higher than 0.47. Such values are indicative of relatively high potential of photosynthetic activity of phytoplankton. Abundance and species diversity were higher than those in the dry season (March-April).

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.

Working on a baseline for the Kongsfjorden food web: production and properties of macroalgal particulate organic matter (POM), supplementary material

Macroalgae, in particular kelps, produce a large amount of biomass in Kongsfjorden, which is to a great extent released into the water in an annual cycle. As an example, the brown alga Alaria esculenta loses its blade gradually, 3 ± 0.8 % of the blade area per day (August 2012), thereby adding to the pool of particulate organic matter (POM) in the fjord. Upon release small thallus pieces are "aging" in that they are prone to leaching and serving as substrate for microorganisms, thus turning into palatable food for suspension and bottom feeders. In order to define a macroalgal baseline for the Kongsfjorden food web, stable isotopes d14C and d15N were measured in individuals of A. esculenta, Saccharina latissima and Laminaria digitata directly sampled after collection and in artificially produced POM (aPOM) of A. esculenta that was allowed to age under experimental conditions. In aPOM from this species sampled in August 2012 the C/N ratios decreased between d1 and d8 of a 14-day culture period in parallel to the fading photosynthetic activity of the algal fragments as demonstrated by use of an Imaging-PAM. Microscopic observations of the aPOM in August 2012 and 2013 revealed the frequent occurrence of small brown algal endo- and epiphytes. First feeding experiments with Mysis oculata (Mysids) and Hiatella arctica (Bivalves) showed that these species can ingest macroalgal POM. The importance of kelp-derived POM for the food web is subject of the current research.

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.

Seawater carbonate chemistry and concentration boundary layers around complex assemblages of macroalgae in a laboratory experiment

Metabolic processes have the potential to modulate the effects of ocean acidification (OA) in nearshore macroalgal beds. We investigated whether natural mixed assemblages of the articulate coralline macroalgae Arthrocardia corymbosa and understory crustose coralline algae (CCA) altered pH and O2 concentrations within and immediately above their canopies. In a unidirectional flume, we tested the effect of water velocity (0-0.1 m/s), bulk seawater pH (ambient pH 8.05, and pH 7.65), and irradiance (photosynthetically saturating light and darkness) on pH and O2 concentration gradients, and the derived concentration boundary layer (CBL) thickness. At bulk seawater pH 7.65 and slow velocities (0 and 0.015 m/s), pH at the CCA surface increased to 7.90-8.00 in the light. Although these manipulations were short term, this indicates a potential daytime buffering capacity that could alleviate the effects of OA. Photosynthetic activity also increased O2 concentrations at the surface of the CCA. However, this moderating capacity was flow dependent; the CBL thickness decreased from an average of 26.8 mm from the CCA surface at 0.015 m/s to 4.1 mm at 0.04 m/s. The reverse trends occurred in the dark, with respiration causing pH and O2 concentrations to decrease at the CCA surface. At all flow velocities the CBL thicknesses (up to 68 mm) were much greater than those previously published, indicating that the presence of canopies can alter the CBL substantially. In situ, the height of macroalgal canopies can be an order of magnitude larger than those used here, indicating that the degree of buffering to OA will be context dependent.

Impact of temperature and species interaction on filamentous cyanobacteria may be more important than salinity and increased pCO2 levels

A future business-as-usual scenario (A1FI) was tested on two bloom-forming cyanobacteria of the Baltic Proper, Nodularia spumigena and Aphanizomenon sp., growing separately and together. The projected scenario was tested in two laboratory experiments where (a) interactive effects of increased temperature and decreased salinity and (b) interactive effects of increased temperature and elevated levels of pCO2 were tested. Increased temperature, from 12 to 16 °C, had a positive effect on the biovolume and photosynthetic activity (F v/F m) of both species. Compared when growing separately, the biovolume of each species was lower when grown together. Decreased salinity, from 7 to 4, and elevated levels of pCO2, from 380 to 960 ppm, had no effect on the biovolume, but on F v/F m of N. spumigena with higher F v/F m in salinity 7. Our results suggest that the projected A1FI scenario might be beneficial for the two species dominating the extensive summer blooms in the Baltic Proper. However, our results further stress the importance of studying interactions between species.

Phytoplankton composition and biomass and its fluorescence parameters in waters of the Nha Trang Bay, South China Sea

Species composition and abundance of phytoplankton and chlorophyll concentration were measured at three horizons of 9 stations in the Nha Trang Bay of the South China Sea in March 1998. Vertical distribution of fluorescence parameters, temperature and irradiance were measured in the 0-18 m layer of the water column at 21 stations. It was shown that according to biomass (B) and chlorophyll concentration (Chl) the Bay is mezotrophic. B and Chl in the water column increased seaward. Mean values of Chl in the southern part of the Bay exceeded those in northern part. Mean values of B were similar. B and Chl in the bottom layer exceeded ones in the upper layer. Diatoms dominated in species diversity and abundance. Diatom Guinardia striata made the main contribution to phytoplankton biomass. Similarity of phytoplankton was high. In the upper layer phytoplankton was photoinhibited during the most part of the light period, but at the bottom photosynthetic activity was high. Water column B varied in an order of magnitude during the daily cycle mainly because of B variations in the bottom layer due to tide flow.

Direct effects of increased CO2 concentrations in seawater on the net primary production of charophytes in a shallow, coastal, brackish-water ecosystem

Charophytes are found in fresh and brackish waters across the globe and play key roles in coastal ecosystems. However, their response to increasing CO2 is not well understood. The aim of the study was to detect the effects of elevated CO2 on the physiology of charophyte species growing in the brackish Baltic Sea by measuring net primary production. Mesocosm experiments were conducted in the Kõiguste Bay (N Gulf of Riga) during the field season of 2012. Separate mesocosms were maintained at different pCO2 levels: 2000, 1000 and 200 µatm. The experiments were carried out with three species of charophytes: Chara aspera, C. tomentosa and C. horrida. The short-term photosynthetic responses of charophytes to different treatments were measured by the oxygen method. The results show that elevated CO2 levels in brackish water may enhance the photosynthetic activity of charophyte species and suggest that increasing CO2 in the Baltic Sea could have implications for interspecific competition and community structure in a future high CO2 world.

Seagrass ecosystem response to long-term high CO2 in a Mediterranean volcanic vent

We examined the long-term effect of naturally acidified water on a Cymodocea nodosa meadow growing at a shallow volcanic CO2 vent in Vulcano Island (Italy). Seagrass and adjacent unvegetated habitats growing at a low pH station (pH = 7.65 ± 0.02) were compared with corresponding habitats at a control station (pH = 8.01 ± 0.01). Density and biomass showed a clear decreasing trend at the low pH station and the below- to above-ground biomass ratio was more than 10 times lower compared to the control. C content and delta 13C of leaves and epiphytes were significantly lower at the low pH station. Photosynthetic activity of C. nodosa was stimulated by low pH as seen by the significant increase in Chla content of leaves, maximum electron transport rate and compensation irradiance. Seagrass community metabolism was intense at the low pH station, with significantly higher net community production, respiration and gross primary production than the control community, whereas metabolism of the unvegetated community did not differ between stations. Productivity was promoted by the low pH, but this was not translated into biomass, probably due to nutrient limitation, grazing or poor environmental conditions. The results indicate that seagrass response in naturally acidified conditions is dependable upon species and geochemical characteristics of the site and highlight the need for a better understanding of complex interactions in these environments.

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.

(Table 1, 2) Foraminifera biometric data, and boron isotopic composition of Globigerinoides sacculifer of sediments from the Ontong-Java Plateau

Sediment samples from the Ontong-Java Plateau in the Pacific and the 90° east ridge in the Indian Ocean were used to investigate whether shell size and early diagenesis affect d11B of the symbiont-bearing planktonic foraminifer Globigerinoides sacculifer. In pristine shells from both study locations we found a systematic increase of d11B and Mg/Ca with shell size. Shells in the sieve size class 515-865 µm revealed d11B values +2.1 to +2.3 per mil higher than shells in the 250-380 µm class. This pattern is most likely due to differences in symbiont photosynthetic activity and its integrated effect on the pH of the foraminiferal microenvironment. We therefore suggest smaller individuals must live at approximately 50-100 m water depth where ambient light levels are lower. Using the empirical calibration curve for d11B in G. sacculifer, only shells larger than 425 µm reflect surface seawater pH. Partial dissolution of shells derived from deeper sediment cores was determined by shell weight analyses and investigation of the shell surface microstructure by scanning electron microscopy. The d11B in partially dissolved shells is up to 2 per mil lower relative to pristine shells of the same size class. In agreement with a relatively higher weight loss in smaller shells, samples from the Ontong-Java Plateau show a more pronounced dissolution effect than larger shells. On the basis of the primary size effect and potential postdepositional dissolution effects, we recommend the use of shells that are visually pristine and, in the case of G. sacculifer, larger than 500 ?m for paleoreconstructions.

Adaptation of a globally important coccolithophore to ocean warming and acidification

Regulating intracellular pH (pHi) is critical for optimising the metabolic activity of corals, yet mechanisms involved in pH regulation and the buffering capacity within coral cells are not well understood. Our study investigated how the presence of symbiotic dinoflagellates affects the response of pHi to pCO2-driven seawater acidification in cells isolated from Pocillopora damicornis. Using the fluorescent dye BCECF-AM, in conjunction with confocal microscopy, we simultaneously characterised the response of pHi in host coral cells and their dinoflagellate symbionts, in symbiotic and non-symbiotic states under saturating light, with and without the photosynthetic inhibitor DCMU. Each treatment was run under control (pH 7.8) and CO2 acidified seawater conditions (decreasing pH from 7.8 - 6.8). After two hours of CO2 addition, by which time the external pH (pHe) had declined to 6.8, the dinoflagellate symbionts had increased their pHi by 0.5 pH units above control levels. In contrast, in both symbiotic and non-symbiotic host coral cells, 15 min of CO2 addition (0.2 pH unit drop in pHe) led to cytoplasmic acidosis equivalent to 0.4 pH units. Despite further seawater acidification over the duration of the experiment, the pHi of non-symbiotic coral cells did not change, though in host cells containing a symbiont cell the pHi recovered to control levels. This recovery was negated when cells were incubated with DCMU. Our results reveal that photosynthetic activity of the endosymbiont is tightly coupled with the ability of the host cell to recover from cellular acidosis after exposure to high CO2 / low pH.

Presence of competitors influences photosynthesis, but not growth, of the hard coral Porites cylindrica at elevated seawater CO2

Changes in environmental conditions, such as those caused by elevated carbon dioxide (CO2), potentially alter the outcome of competitive interactions between species. This study aimed to understand how elevated CO2 could influence competitive interactions between hard and soft corals, by investigating growth and photosynthetic activity of Porites cylindrica (a hard coral) under elevated CO2 and in the presence of another hard coral and two soft coral competitors. Corals were collected from reefs around Orpheus and Pelorus Islands on the Great Barrier Reef, Australia. They were then exposed to elevated pCO2 for 4 weeks with two CO2 treatments: intermediate (pCO2 648) and high (pCO2 1003) compared with a control (unmanipulated seawater) treatment (pCO2 358). Porites cylindrica growth did not vary among pCO2 treatments, regardless of the presence and type of competitors, nor was the growth of another hard coral species, Acropora cerealis, affected by pCO2 treatment. Photosynthetic rates of P. cylindrica were sensitive to variations in pCO2, and varied between the side of the fragment facing the competitors vs. the side facing away from the competitor. However, variation in photosynthetic rates depended on pCO2 treatment, competitor identity, and whether the photosynthetic yields were measured as maximum or effective photosynthetic yield. This study suggests that elevated CO2 may impair photosynthetic activity, but not growth, of a hard coral under competition and confirms the hypothesis that soft corals are generally resistant to elevated CO2. Overall, our results indicate that shifts in the species composition in coral communities as a result of elevated CO2 could be more strongly related to the individual tolerance of different species rather than a result of competitive interactions between species.