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      Growth, ammonium metabolism, and photosynthetic properties of Ulva australis (Chlorophyta) under decreasing pH and ammonium enrichment

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          Abstract

          The responses of macroalgae to ocean acidification could be altered by availability of macronutrients, such as ammonium (NH 4 +). This study determined how the opportunistic macroalga, Ulva australis responded to simultaneous changes in decreasing pH and NH 4 + enrichment. This was investigated in a week-long growth experiment across a range of predicted future pHs with ambient and enriched NH 4 + treatments followed by measurements of relative growth rates (RGR), NH 4 + uptake rates and pools, total chlorophyll, and tissue carbon and nitrogen content. Rapid light curves (RLCs) were used to measure the maximum relative electron transport rate (rETR max) and maximum quantum yield of photosystem II (PSII) photochemistry (F v/F m). Photosynthetic capacity was derived from the RLCs and included the efficiency of light harvesting (α), slope of photoinhibition (β), and the light saturation point (E k). The results showed that NH 4 + enrichment did not modify the effects of pH on RGRs, NH 4 + uptake rates and pools, total chlorophyll, rETR max, α, β, F v/F m, tissue C and N, and the C:N ratio. However, E k was differentially affected by pH under different NH 4 + treatments. E k increased with decreasing pH in the ambient NH 4 + treatment, but not in the enriched NH 4 + treatment. NH 4 + enrichment increased RGRs, NH 4 + pools, total chlorophyll, rETR max, α, β, F v/F m, and tissue N, and decreased NH 4 + uptake rates and the C:N ratio. Decreased pH increased total chlorophyll content, rETR max, F v/F m, and tissue N content, and decreased the C:N ratio. Therefore, the results indicate that U. australis growth is increased with NH 4 + enrichment and not with decreasing pH. While decreasing pH influenced the carbon and nitrogen metabolisms of U. australis, it did not result in changes in growth.

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          Ocean acidification: the other CO2 problem.

          Rising atmospheric carbon dioxide (CO2), primarily from human fossil fuel combustion, reduces ocean pH and causes wholesale shifts in seawater carbonate chemistry. The process of ocean acidification is well documented in field data, and the rate will accelerate over this century unless future CO2 emissions are curbed dramatically. Acidification alters seawater chemical speciation and biogeochemical cycles of many elements and compounds. One well-known effect is the lowering of calcium carbonate saturation states, which impacts shell-forming marine organisms from plankton to benthic molluscs, echinoderms, and corals. Many calcifying species exhibit reduced calcification and growth rates in laboratory experiments under high-CO2 conditions. Ocean acidification also causes an increase in carbon fixation rates in some photosynthetic organisms (both calcifying and noncalcifying). The potential for marine organisms to adapt to increasing CO2 and broader implications for ocean ecosystems are not well known; both are high priorities for future research. Although ocean pH has varied in the geological past, paleo-events may be only imperfect analogs to current conditions.
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            Impacts of ocean acidification on marine organisms: quantifying sensitivities and interaction with warming

            Ocean acidification represents a threat to marine species worldwide, and forecasting the ecological impacts of acidification is a high priority for science, management, and policy. As research on the topic expands at an exponential rate, a comprehensive understanding of the variability in organisms' responses and corresponding levels of certainty is necessary to forecast the ecological effects. Here, we perform the most comprehensive meta-analysis to date by synthesizing the results of 228 studies examining biological responses to ocean acidification. The results reveal decreased survival, calcification, growth, development and abundance in response to acidification when the broad range of marine organisms is pooled together. However, the magnitude of these responses varies among taxonomic groups, suggesting there is some predictable trait-based variation in sensitivity, despite the investigation of approximately 100 new species in recent research. The results also reveal an enhanced sensitivity of mollusk larvae, but suggest that an enhanced sensitivity of early life history stages is not universal across all taxonomic groups. In addition, the variability in species' responses is enhanced when they are exposed to acidification in multi-species assemblages, suggesting that it is important to consider indirect effects and exercise caution when forecasting abundance patterns from single-species laboratory experiments. Furthermore, the results suggest that other factors, such as nutritional status or source population, could cause substantial variation in organisms' responses. Last, the results highlight a trend towards enhanced sensitivity to acidification when taxa are concurrently exposed to elevated seawater temperature.
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              CO2 concentrating mechanisms in algae: mechanisms, environmental modulation, and evolution.

              The evolution of organisms capable of oxygenic photosynthesis paralleled a long-term reduction in atmospheric CO2 and the increase in O2. Consequently, the competition between O2 and CO2 for the active sites of RUBISCO became more and more restrictive to the rate of photosynthesis. In coping with this situation, many algae and some higher plants acquired mechanisms that use energy to increase the CO2 concentrations (CO2 concentrating mechanisms, CCMs) in the proximity of RUBISCO. A number of CCM variants are now found among the different groups of algae. Modulating the CCMs may be crucial in the energetic and nutritional budgets of a cell, and a multitude of environmental factors can exert regulatory effects on the expression of the CCM components. We discuss the diversity of CCMs, their evolutionary origins, and the role of the environment in CCM modulation.
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                Author and article information

                Contributors
                Role: ConceptualizationRole: Data curationRole: Formal analysisRole: Funding acquisitionRole: InvestigationRole: MethodologyRole: Project administrationRole: VisualizationRole: Writing – original draftRole: Writing – review & editing
                Role: ConceptualizationRole: InvestigationRole: MethodologyRole: Writing – review & editing
                Role: InvestigationRole: MethodologyRole: Writing – review & editing
                Role: InvestigationRole: MethodologyRole: Writing – review & editing
                Role: ConceptualizationRole: MethodologyRole: ResourcesRole: SoftwareRole: Writing – review & editing
                Role: ResourcesRole: Writing – review & editing
                Role: ConceptualizationRole: Funding acquisitionRole: MethodologyRole: Project administrationRole: ResourcesRole: SoftwareRole: SupervisionRole: Writing – review & editing
                Role: ConceptualizationRole: MethodologyRole: SupervisionRole: Writing – review & editing
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                27 November 2017
                2017
                : 12
                : 11
                : e0188389
                Affiliations
                [1 ] Department of Biology, California State University at Northridge, Northridge, California, United States of America
                [2 ] Institute for Marine and Antarctic Studies, University of Tasmania, Hobart, Tasmania, Australia
                [3 ] GEOMAR Helmholtz Centre for Ocean Research, Kiel, Germany
                [4 ] Department of Chemistry, University of Otago, Dunedin, New Zealand
                [5 ] CSIRO, Oceans and Atmosphere, Hobart, Tasmania, Australia
                The University of Hong Kong, HONG KONG
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                [¤a]

                Current address: School of Marine and Atmospheric Sciences, Stony Brook University, Stony Brook, New York, United States of America

                [¤b]

                Current address: Centro i~mar, Universidad de Los Lagos, Puerto Montt, Chile

                [¤c]

                Current address: Instituto de Fomento Pesquero, Puerto Montt, Chile

                Author information
                http://orcid.org/0000-0003-4445-4621
                Article
                PONE-D-17-13507
                10.1371/journal.pone.0188389
                5703455
                29176815
                46168a3c-86ea-4045-abc8-641e6e7de2ac
                © 2017 Reidenbach et al

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 6 April 2017
                : 6 November 2017
                Page count
                Figures: 6, Tables: 2, Pages: 20
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/100000001, National Science Foundation;
                Award ID: 1515267
                Award Recipient :
                This work was supported by National Science Foundation grant: OISE 1515267 (URL: https://www.nsf.gov/awardsearch/showAward?AWD_ID=1515267) (Author: LBR). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Ecology and Environmental Sciences
                Aquatic Environments
                Marine Environments
                Sea Water
                Earth Sciences
                Marine and Aquatic Sciences
                Aquatic Environments
                Marine Environments
                Sea Water
                Biology and Life Sciences
                Organisms
                Eukaryota
                Plants
                Algae
                Biology and Life Sciences
                Cell Biology
                Cellular Structures and Organelles
                Chloroplasts
                Chlorophyll
                Biology and Life Sciences
                Cell Biology
                Plant Cell Biology
                Chloroplasts
                Chlorophyll
                Biology and Life Sciences
                Plant Science
                Plant Cell Biology
                Chloroplasts
                Chlorophyll
                Biology and Life Sciences
                Cell Biology
                Cellular Types
                Plant Cells
                Chloroplasts
                Chlorophyll
                Biology and Life Sciences
                Cell Biology
                Plant Cell Biology
                Plant Cells
                Chloroplasts
                Chlorophyll
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                Plant Cells
                Chloroplasts
                Chlorophyll
                Physical Sciences
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                Pigments
                Organic Pigments
                Chlorophyll
                Physical Sciences
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                Biology and Life Sciences
                Plant Science
                Plant Biochemistry
                Photosynthesis
                Earth Sciences
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                Oceanography
                Ocean Acidification
                Physical Sciences
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                Particle Physics
                Elementary Particles
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                Custom metadata
                All data files are available from the CSUN ScholarWorks Open Access Repository (SOAR) ( http://hdl.handle.net/10211.3/195049).

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