Understanding coralline algal responses to ocean acidification: Meta-analysis and synthesis.

Ocean acidification (OA) is a major threat to the persistence of biogenic reefs throughout the world's ocean. Coralline algae are comprised of high magnesium calcite and have long been considered one of the most susceptible taxa to the negative impacts of OA. We summarize these impacts and explore the causes of variability in coralline algal responses using a review/qualitative assessment of all relevant literature, meta-analysis, quantitative assessment of critical responses, and a discussion of physiological mechanisms and directions for future research. We find that most coralline algae experienced reduced abundance, calcification rates, recruitment rates, and declines in pH within the site of calcification in laboratory experiments simulating OA or at naturally elevated CO2 sites. There were no other consistent physiological responses of coralline algae to simulated OA (e.g., photo-physiology, mineralogy, and survival). Calcification/growth was the most frequently measured parameters in coralline algal OA research, and our meta-analyses revealed greater declines in seawater pH were associated with significant decreases in calcification in adults and similar but nonsignificant trends for juveniles. Adults from the family Mesophyllumaceae also tended to be more robust to OA, though there was insufficient data to test similar trends for juveniles. OA was the dominant driver in the majority of laboratory experiments where other local or global drivers were assessed. The interaction between OA and any other single driver was often additive, though factors that changed pH at the surface of coralline algae (light, water motion, epiphytes) acted antagonistically or synergistically with OA more than any other drivers. With advances in experimental design and methodological techniques, we now understand that the physiology of coralline algal calcification largely dictates their responses to OA. However, significant challenges still remain, including improving the geographic and life-history spread of research effort and a need for holistic assessments of physiology.

Ocean acidification locks algal communities in a species-poor early successional stage.

Long-term exposure to CO2 -enriched waters can considerably alter marine biological community development, often resulting in simplified systems dominated by turf algae that possess reduced biodiversity and low ecological complexity. Current understanding of the underlying processes by which ocean acidification alters biological community development and stability remains limited, making the management of such shifts problematic. Here, we deployed recruitment tiles in reference (pHT 8.137 ± 0.056 SD) and CO2 -enriched conditions (pHT 7.788 ± 0.105 SD) at a volcanic CO2 seep in Japan to assess the underlying processes and patterns of algal community development. We assessed (i) algal community succession in two different seasons (Cooler months: January-July, and warmer months: July-January), (ii) the effects of initial community composition on subsequent community succession (by reciprocally transplanting preestablished communities for a further 6 months), and (iii) the community production of resulting communities, to assess how their functioning was altered (following 12 months recruitment). Settlement tiles became dominated by turf algae under CO2 -enrichment and had lower biomass, diversity and complexity, a pattern consistent across seasons. This locked the community in a species-poor early successional stage. In terms of community functioning, the elevated pCO2 community had greater net community production, but this did not result in increased algal community cover, biomass, biodiversity or structural complexity. Taken together, this shows that both new and established communities become simplified by rising CO2 levels. Our transplant of preestablished communities from enriched CO2 to reference conditions demonstrated their high resilience, since they became indistinguishable from communities maintained entirely in reference conditions. This shows that meaningful reductions in pCO2 can enable the recovery of algal communities. By understanding the ecological processes responsible for driving shifts in community composition, we can better assess how communities are likely to be altered by ocean acidification.

Simplification, not "tropicalization", of temperate marine ecosystems under ocean warming and acidification.

Ocean warming is altering the biogeographical distribution of marine organisms. In the tropics, rising sea surface temperatures are restructuring coral reef communities with sensitive species being lost. At the biogeographical divide between temperate and tropical communities, warming is causing macroalgal forest loss and the spread of tropical corals, fishes and other species, termed "tropicalization". A lack of field research into the combined effects of warming and ocean acidification means there is a gap in our ability to understand and plan for changes in coastal ecosystems. Here, we focus on the tropicalization trajectory of temperate marine ecosystems becoming coral-dominated systems. We conducted field surveys and in situ transplants at natural analogues for present and future conditions under (i) ocean warming and (ii) both ocean warming and acidification at a transition zone between kelp and coral-dominated ecosystems. We show that increased herbivory by warm-water fishes exacerbates kelp forest loss and that ocean acidification negates any benefits of warming for range extending tropical corals growth and physiology at temperate latitudes. Our data show that, as the combined effects of ocean acidification and warming ratchet up, marine coastal ecosystems lose kelp forests but do not gain scleractinian corals. Ocean acidification plus warming leads to overall habitat loss and a shift to simple turf-dominated ecosystems, rather than the complex coral-dominated tropicalized systems often seen with warming alone. Simplification of marine habitats by increased CO2 levels cascades through the ecosystem and could have severe consequences for the provision of goods and services.

Simple and convenient preconcentration procedure for the isotopic analysis of uranium in seawater.

The demand for monitoring anthropogenic U isotopes, 236U and 233U, in seawater will continue to increase due to radioecological issues and the need for tools for environmental dynamics research. In response to this growing demand, herein, a novel and simple method was developed for the collection of U isotopes in seawater, both in the laboratory and field, using a fabric-like amidoxime adsorbent. The results from the adsorption studies showed that the optimum conditions for processing seawater in a glass beaker were as follows: seawater pH 4, amidoxime adsorbent 0.20 mmol per 500 g seawater and an adsorption time of 9 hours. Alternatively, when using a closed polyethylene container in experiments on-board a ship and using the same ratio of adsorbent to seawater as in the beaker experiment in the laboratory, the optimum conditions were as follows: seawater pH 8 and an adsorption time of 24 hours. Under the above-mentioned conditions, more than 95% of the U underwent adsorption in both the beaker and the polyethylene container experiments. In the case of analyte desorption, more than 80% of U in seawater was recovered using 2-3 mol dm-3 HCl or HNO3 as the eluent. Thus, it was concluded that the amidoxime adsorbent can serve as a simple and effective pre-concentration method for the ultra-trace monitoring of U isotopes in seawater.

Ocean acidification increases the impact of typhoons on algal communities.

Long-term environmental change, sudden pulses of extreme perturbation, or a combination of both can trigger regime shifts by changing the processes and feedbacks which determine community assembly, structure, and function, altering the state of ecosystems. Our understanding of the mechanisms that stabilise against regime shifts or lock communities into altered states is limited, yet also critical to anticipating future states, preventing regime shifts, and reversing unwanted state change. Ocean acidification contributes to the restructuring and simplification of algal systems, however the mechanisms through which this occurs and whether additional drivers are involved requires further study. Using monthly surveys over three years at a shallow-water volcanic seep we examined how the composition of algal communities change seasonally and following periods of significant physical disturbance by typhoons at three levels of ocean acidification (equivalent to means of contemporary ∼350 and future ∼500 and 900 µatm pCO2). Sites exposed to acidification were increasingly monopolised by structurally simple, fast-growing turf algae, and were clearly different to structurally complex macrophyte-dominated reference sites. The distinct contemporary and acidified community states were stabilised and maintained at their respective sites by different mechanisms following seasonal typhoon disturbance. Macroalgal-dominated sites were resistant to typhoon damage. In contrast, significant losses of algal biomass represented a near total ecosystem reset by typhoons for the turf-dominated communities at the elevated pCO2 sites (i.e. negligible resistance). A combination of disturbance and subsequent turf recovery maintained the same simplified state between years (elevated CO2 levels promote turf growth following algal removal, inhibiting macroalgal recruitment). Thus, ocean acidification may promote shifts in algal systems towards degraded ecosystem states, and short-term disturbances which reset successional trajectories may 'lock-in' these alternative states of low structural and functional diversity.

Tetracycline resistance genes persist at aquaculture farms in the absence of selection pressure.

The prophylactic and therapeutic use of tetracyclines in aquaculture has been shown to contribute to the spread of tetracycline resistance in the environment. In this work, the prevalence of four different tetracycline-resistance genes, tetA, tetC, tetH, and tetM, in sediments from four aquaculture farms and their surroundings in the Baltic Sea was monitored by quantitative polymerase chain reaction (qPCR). The presence of three additional tetracycline-resistance genes (tetE, tetG, and tetW) was studied qualitatively by standard PCR, and the amount of bioavailable tetracyclines and total amounts of tetracycline and oxytetracycline in samples were also measured. None of the farms were using tetracycline at the time of the sampling and one of the farms had stopped all antibiotic use six years prior to the first sampling. Two of the farms were sampled over four successive summers and two were sampled once. Our results showed greater copy numbers of tetA, tetC, tetH, and tetM at the farms compared to pristine sites and demonstrated the presence of tetE, tetG, and tetW genes in the sediments under aquaculture farms at most sampling times. However, no resistance genes were found in samples collected 200 m from any of the farms. None of the samples contained therapeutically active concentrations of tetracyclines at any of the sampling times, suggesting that the increase in the prevalence of tetracycline resistance genes is caused by the persistence of these genes in the absence of selection pressure.

Feedback mechanisms stabilise degraded turf algal systems at a CO2 seep site.

Human activities are rapidly changing the structure and function of coastal marine ecosystems. Large-scale replacement of kelp forests and coral reefs with turf algal mats is resulting in homogenous habitats that have less ecological and human value. Ocean acidification has strong potential to substantially favour turf algae growth, which led us to examine the mechanisms that stabilise turf algal states. Here we show that ocean acidification promotes turf algae over corals and macroalgae, mediating new habitat conditions that create stabilising feedback loops (altered physicochemical environment and microbial community, and an inhibition of recruitment) capable of locking turf systems in place. Such feedbacks help explain why degraded coastal habitats persist after being initially pushed past the tipping point by global and local anthropogenic stressors. An understanding of the mechanisms that stabilise degraded coastal habitats can be incorporated into adaptive management to better protect the contribution of coastal systems to human wellbeing.

Changes in fish communities due to benthic habitat shifts under ocean acidification conditions.

Ocean acidification will likely change the structure and function of coastal marine ecosystems over coming decades. Volcanic carbon dioxide seeps generate dissolved CO2 and pH gradients that provide realistic insights into the direction and magnitude of these changes. Here, we used fish and benthic community surveys to assess the spatio-temporal dynamics of fish community properties off CO2 seeps in Japan. Adding to previous evidence from ocean acidification ecosystem studies conducted elsewhere, our findings documented shifts from calcified to non-calcified habitats with reduced benthic complexity. In addition, we found that such habitat transition led to decreased diversity of associated fish and to selection of those fish species better adapted to simplified ecosystems dominated by algae. Our data suggest that near-future projected ocean acidification levels will oppose the ongoing range expansion of coral reef-associated fish due to global warming.

Ocean acidification drives community shifts towards simplified non-calcified habitats in a subtropical-temperate transition zone.

Rising atmospheric concentrations of carbon dioxide are causing surface seawater pH and carbonate ion concentrations to fall in a process known as ocean acidification. To assess the likely ecological effects of ocean acidification we compared intertidal and subtidal marine communities at increasing levels of pCO2 at recently discovered volcanic seeps off the Pacific coast of Japan (34° N). This study region is of particular interest for ocean acidification research as it has naturally low levels of surface seawater pCO2 (280-320 µatm) and is located at a transition zone between temperate and sub-tropical communities. We provide the first assessment of ocean acidification effects at a biogeographic boundary. Marine communities exposed to mean levels of pCO2 predicted by 2050 experienced periods of low aragonite saturation and high dissolved inorganic carbon. These two factors combined to cause marked community shifts and a major decline in biodiversity, including the loss of key habitat-forming species, with even more extreme community changes expected by 2100. Our results provide empirical evidence that near-future levels of pCO2 shift sub-tropical ecosystems from carbonate to fleshy algal dominated systems, accompanied by biodiversity loss and major simplification of the ecosystem.

The effect of zinc on aquatic microbial ecosystems and the degradation of dissolved organic matter.

We developed a simple aquatic microbial ecosystem model in order to examine potential effects of zinc on microorganisms and the related degradation of dissolved organic matter (DOM). The model is a combination of both a traditional food chain and a microbial loop. The traditional food chain is mainly composed of phytoplankton and zooplankton, whilst the microbial loop is composed of DOM, bacteria and bacterivorous protozoa. We incorporated the suppressive effect of zinc on the bacterial uptake of DOM and assessed the steady state responses of the model for various zinc concentrations. The analytical and numerical results of the model implied that either zooplankton or bacterivorous protozoa might be the most vulnerable group to excessive zinc load than bacteria, depending on the grazing preference of zooplankton between phytoplankton and bacterivorous protozoa. The sensitivity analyses supported that the microbial loop solely is more sensitive to zinc than the coupled system combining both the traditional food chain and the microbial loop.