Resilience of Emiliania huxleyi to future changes in subantarctic waters.

Lower pH and elevated temperature alter phytoplankton growth and biomass in short-term incubations, but longer-term responses and adaptation potential are less well-studied. To determine the future of the coccolithophore Emiliania huxleyi, a mixed genotype culture from subantarctic water was incubated for 720 days under present-day temperature and pH, and also projected future conditions by the year 2100. The future population exhibited a higher growth rate relative to present-day cells transferred to future conditions after 309 days, indicating adaptation or genotype selection; this was reflected by an increase in optimum growth temperature of ~2.5°C by the end of the experiment. Following transfer to opposing conditions in short-term cross-over incubations, cell volume responded rapidly, within eight generations, confirming trait plasticity. The changes in growth rate and cell volume were larger than reported in previous single stressor relationships and incubations, suggesting synergistic or additive effects of combined elevated temperature and lower pH and highlighting the importance of long-term multiple stressor experiments. At the end of the incubation there were no significant differences in cellular composition (particulate organic content and chlorophyll a), or primary production between present-day and future populations. Conversely, two independent methods showed a 50% decrease in both particulate inorganic carbon and calcification rate, consistent with the decrease in cell volume, in the future population. The observed plasticity and adaptive capacity of E. huxleyi indicate resilience to future conditions in subantarctic waters, although changes in cell volume and carbonate may alter grazing loss and cell ballast, so influencing carbon export to the deep ocean.

Evidence of nitrate-based nighttime atmospheric nucleation driven by marine microorganisms in the South Pacific.

Our understanding of ocean-cloud interactions and their effect on climate lacks insight into a key pathway: do biogenic marine emissions form new particles in the open ocean atmosphere? Using measurements collected in ship-borne air-sea interface tanks deployed in the Southwestern Pacific Ocean, we identified new particle formation (NPF) during nighttime that was related to plankton community composition. We show that nitrate ions are the only species for which abundance could support NPF rates in our semicontrolled experiments. Nitrate ions also prevailed in the natural pristine marine atmosphere and were elevated under higher sub-10 nm particle concentrations. We hypothesize that these nucleation events were fueled by complex, short-term biogeochemical cycling involving the microbial loop. These findings suggest a new perspective with a previously unidentified role of nitrate of marine biogeochemical origin in aerosol nucleation.

Forensic carbon accounting: Assessing the role of seaweeds for carbon sequestration.

Carbon sequestration is defined as the secure storage of carbon-containing molecules for >100 years, and in the context of carbon dioxide removal for climate mitigation, the origin of this CO2 is from the atmosphere. On land, trees globally sequester substantial amounts of carbon in woody biomass, and an analogous role for seaweeds in ocean carbon sequestration has been suggested. The purposeful expansion of natural seaweed beds and aquaculture systems, including into the open ocean (ocean afforestation), has been proposed as a method of increasing carbon sequestration and use in carbon trading and offset schemes. However, to verify whether CO2 fixed by seaweeds through photosynthesis leads to carbon sequestration is extremely complex in the marine environment compared to terrestrial systems, because of the need to jointly consider: the comparatively rapid turnover of seaweed biomass, tracing the fate of carbon via particulate and dissolved organic carbon pathways in dynamic coastal waters, and the key role of atmosphere-ocean CO2 exchange. We propose a Forensic Carbon Accounting approach, in which a thorough analysis of carbon flows between the atmosphere and ocean, and into and out of seaweeds would be undertaken, for assessing the magnitude of CO2 removal and robust attribution of carbon sequestration to seaweeds.

Surface ocean microbiota determine cloud precursors.

One pathway by which the oceans influence climate is via the emission of sea spray that may subsequently influence cloud properties. Sea spray emissions are known to be dependent on atmospheric and oceanic physicochemical parameters, but the potential role of ocean biology on sea spray fluxes remains poorly characterized. Here we show a consistent significant relationship between seawater nanophytoplankton cell abundances and sea-spray derived Cloud Condensation Nuclei (CCN) number fluxes, generated using water from three different oceanic regions. This sensitivity of CCN number fluxes to ocean biology is currently unaccounted for in climate models yet our measurements indicate that it influences fluxes by more than one order of magnitude over the range of phytoplankton investigated.

The impacts of ocean acidification on marine trace gases and the implications for atmospheric chemistry and climate.

Surface ocean biogeochemistry and photochemistry regulate ocean-atmosphere fluxes of trace gases critical for Earth's atmospheric chemistry and climate. The oceanic processes governing these fluxes are often sensitive to the changes in ocean pH (or pCO2) accompanying ocean acidification (OA), with potential for future climate feedbacks. Here, we review current understanding (from observational, experimental and model studies) on the impact of OA on marine sources of key climate-active trace gases, including dimethyl sulfide (DMS), nitrous oxide (N2O), ammonia and halocarbons. We focus on DMS, for which available information is considerably greater than for other trace gases. We highlight OA-sensitive regions such as polar oceans and upwelling systems, and discuss the combined effect of multiple climate stressors (ocean warming and deoxygenation) on trace gas fluxes. To unravel the biological mechanisms responsible for trace gas production, and to detect adaptation, we propose combining process rate measurements of trace gases with longer term experiments using both model organisms in the laboratory and natural planktonic communities in the field. Future ocean observations of trace gases should be routinely accompanied by measurements of two components of the carbonate system to improve our understanding of how in situ carbonate chemistry influences trace gas production. Together, this will lead to improvements in current process model capabilities and more reliable predictions of future global marine trace gas fluxes.

Effects of multiple drivers of ocean global change on the physiology and functional gene expression of the coccolithophore Emiliania huxleyi.

Ongoing ocean global change due to anthropogenic activities is causing multiple chemical and physical seawater properties to change simultaneously, which may affect the physiology of marine phytoplankton. The coccolithophore Emiliania huxleyi is a model species often employed in the study of the marine carbon cycle. The effect of ocean acidification (OA) on coccolithophore calcification has been extensively studied; however, physiological responses to multiple environmental drivers are still largely unknown. Here we examined two-way and multiple driver effects of OA and other key environmental drivers-nitrate, phosphate, irradiance, and temperature-on the growth, photosynthetic, and calcification rates, and the elemental composition of E. huxleyi. In addition, changes in functional gene expression were examined to understand the molecular mechanisms underpinning the physiological responses. The single driver manipulation experiments suggest decreased nitrate supply being the most important driver regulating E. huxleyi physiology, by significantly reducing the growth, photosynthetic, and calcification rates. In addition, the interaction of OA and decreased nitrate supply (projected for year 2100) had more negative synergistic effects on E. huxleyi physiology than all other two-way factorial manipulations, suggesting a linkage between the single dominant driver (nitrate) effects and interactive effects with other drivers. Simultaneous manipulation of all five environmental drivers to the conditions of the projected year 2100 had the largest negative effects on most of the physiological metrics. Furthermore, functional genes associated with inorganic carbon acquisition (RubisCO, AEL1, and δCA) and calcification (CAX3, AEL1, PATP, and NhaA2) were most downregulated by the multiple driver manipulation, revealing linkages between responses of functional gene expression and associated physiological metrics. These findings together indicate that for more holistic projections of coccolithophore responses to future ocean global change, it is necessary to understand the relative importance of environmental drivers both individually (i.e., mechanistic understanding) and interactively (i.e., cumulative effect) on coccolithophore physiology.

Subtle bacterioplankton community responses to elevated CO2 and warming in the oligotrophic South Pacific gyre.

Bacterioplankton play a critical role in primary production, carbon cycling, and nutrient cycling in the oligotrophic ocean. To investigate the effect of elevated CO2 and warming on the composition and function of bacterioplankton communities in oligotrophic waters, we performed two trace-metal clean deck board incubation experiments during the New Zealand GEOTRACES transect of the South Pacific gyre (SPG). High-throughput amplicon sequencing of the 16S rRNA gene revealed that bacterioplankton community composition was distinct between the fringe and ultra-oligotrophic centre of the SPG and changed consistently in response to elevated CO2 at the ultra-oligotrophic centre but not at the mesotrophic fringe of the SPG. The combined effects of elevated CO2 and warming resulted in a high degree of heterogeneity between replicate communities. Community-level protein synthesis rates (3 H-Leucine incorporation) and bacterioplankton abundance were not affected by elevated CO2 alone or in combination with warming at the fringe or ultra-oligotrophic centre of the SPG. These data suggest bacterioplankton community responses to elevated CO2 may be modulated by nutrient regimes in open ocean ecosystems and highlight the need for further investigation in expanding oligotrophic subtropical gyres.

The response of the marine nitrogen cycle to ocean acidification.

Ocean acidification (OA), arising from the influx of anthropogenically generated carbon, poses a massive threat to the ocean ecosystems. Our knowledge of the effects of elevated anthropogenic CO2 in marine waters and its effect on the performance of single species, trophic interactions, and ecosystems is increasing rapidly. However, our understanding of the biogeochemical cycling of nutrients such as nitrogen is less advanced and lacks a comprehensive overview of how these processes may change under OA. We conducted a systematic review and meta-analysis of eight major nitrogen transformation processes incorporating 49 publications to synthesize current scientific understanding of the effect of OA on nitrogen cycling in the future ocean by 2100. The following points were identified by our meta-analysis: (a) Diazotrophic nitrogen fixation is likely enhanced by 29% ± 4% under OA; (b) species- and strain-specific responses of nitrogen fixers to OA were detectable, which may result in alterations in microbial community composition in the future ocean; (c) nitrification processes were reduced by a factor of 29% ± 10%; (d) declines in nitrification rates were not reflected by nitrifier abundance; and (e) contrasting results in unispecific culture experiments versus natural communities were apparent for nitrogen fixation and denitrification. The net effect of the nitrogen cycle process responses also suggests there may be a shift in the relative nitrogen pools, with excess ammonium originating from CO2 -fertilized diazotrophs. This regenerated inorganic nitrogen may recycle in the upper water column increasing the relative importance of the ammonium-fueled regenerated production. However, several feedback mechanisms with other chemical cycles, such as oxygen, and interaction with other climate change stressors may counteract these findings. Finally, our review highlights the shortcomings and gaps in current understanding of the potential changes in nitrogen cycling under future climate and emphasizes the need for further ecosystem studies.

Bacterial abundance, processes and diversity responses to acidification at a coastal CO2 vent.

Shallow CO2 vents are used as natural laboratories to study biological responses to ocean acidification, and so it is important to determine whether pH is the primary driver of bacterial processes and community composition, or whether other variables associated with vent water have a significant influence. Water from a CO2 vent (46 m, Bay of Plenty, New Zealand) was compared to reference water from an upstream control site, and also to control water acidified to the same pH as the vent water. After 84 h, both vent and acidified water exhibited higher potential bulk water and cell-specific glucosidase activity relative to control water, whereas cell-specific protease activities were similar. However, bulk vent water glucosidase activity was double that of the acidified water, as was bacterial secondary production in one experiment, suggesting that pH was not the only factor affecting carbohydrate hydrolysis. In addition, there were significant differences in bacterial community composition in the vent water relative to the control and acidified water after 84 h, including the presence of extremophiles which may influence carbohydrate degradation. This highlights the importance of characterizing microbial processes and community composition in CO2 vent emissions, to confirm that they represent robust analogues for the future acidified ocean.

No stimulation of nitrogen fixation by non-filamentous diazotrophs under elevated CO2 in the South Pacific.

Nitrogen fixation by diazotrophic cyanobacteria is a critical source of new nitrogen to the oligotrophic surface ocean. Research to date indicates that some diazotroph groups may increase nitrogen fixation under elevated pCO2 . To test this in natural plankton communities, four manipulation experiments were carried out during two voyages in the South Pacific (30-35o S). High CO2 treatments, produced using 750 ppmv CO2 to adjust pH to 0.2 below ambient, and 'Greenhouse' treatments (0.2 below ambient pH and ambient temperature +3 °C), were compared with Controls in trace metal clean deckboard incubations in triplicate. No significant change was observed in nitrogen fixation in either the High CO2 or Greenhouse treatments over 5 day incubations. qPCR measurements and optical microscopy determined that the diazotroph community was dominated by Group A unicellular cyanobacteria (UCYN-A), which may account for the difference in response of nitrogen fixation under elevated CO2 to that reported previously for Trichodesmium. This may reflect physiological differences, in that the greater cell surface area:volume of UCYN-A and its lack of metabolic pathways involved in carbon fixation may confer no benefit under elevated CO2 . However, multiple environmental controls may also be a factor, with the low dissolved iron concentrations in oligotrophic surface waters limiting the response to elevated CO2 . If nitrogen fixation by UCYN-A is not stimulated by elevated pCO2 , then future increases in CO2 and warming may alter the regional distribution and dominance of different diazotroph groups, with implications for dissolved iron availability and new nitrogen supply in oligotrophic regions.

Relating carbon monoxide photoproduction to dissolved organic matter functionality.

Aqueous solutions of humic substances (HSs) and pure monomeric aromatics were irradiated to investigate the chemical controls upon carbon monoxide (CO) photoproduction from dissolved organic matter (DOM). HSs were isolated from lakes, rivers, marsh, and ocean. Inclusion of humic, fulvic, hydrophobic organic, and hydrophilic organic acid fractions from these environments provided samples diverse in source and isolation protocol. In spite of these major differences, HS absorption coefficients (a) and photoreactivities (a bleaching and CO production) were strongly dependent upon HS aromaticity (r2 > 0.90; n = 11), implying aromatic moieties are the principal chromophores and photoreactants within HSs, and by extension, DOM. Carbonyl carbon and CO photoproduction were not correlated, implying that carbonyl moieties are not quantitatively important in CO photoproduction. CO photoproduction efficiency of aqueous solutions of monomeric aromatic compounds that are common constituents of organic matter varied with the nature of ring substituents. Specifically, electron donating groups increased, while electron withdrawing groups decreased CO photoproductivity, supporting our conclusion that carbonyl substituents are not quantitatively important in CO photoproduction. Significantly, aromatic CO photoproduction efficiency spanned 3 orders of magnitude, indicating that variations in the CO apparent quantum yields of natural DOM may be related to variations in aromatic DOM substituent group chemistry.

The decline and fate of an iron-induced subarctic phytoplankton bloom.

Iron supply has a key role in stimulating phytoplankton blooms in high-nitrate low-chlorophyll oceanic waters. However, the fate of the carbon fixed by these blooms, and how efficiently it is exported into the ocean's interior, remains largely unknown. Here we report on the decline and fate of an iron-stimulated diatom bloom in the Gulf of Alaska. The bloom terminated on day 18, following the depletion of iron and then silicic acid, after which mixed-layer particulate organic carbon (POC) concentrations declined over six days. Increased particulate silica export via sinking diatoms was recorded in sediment traps at depths between 50 and 125 m from day 21, yet increased POC export was not evident until day 24. Only a small proportion of the mixed-layer POC was intercepted by the traps, with more than half of the mixed-layer POC deficit attributable to bacterial remineralization and mesozooplankton grazing. The depletion of silicic acid and the inefficient transfer of iron-increased POC below the permanent thermocline have major implications both for the biogeochemical interpretation of times of greater iron supply in the geological past, and also for proposed geo-engineering schemes to increase oceanic carbon sequestration.