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1 How Does Ocean Acidification Impact Phytoplankton Productivity And How does ocean acidification impact phytoplankton productivity and community structure? Aliza Ray1 and Joe Vallino2 1Bard College, Annandale-On-Hudson NY 12504 2Marine Biological Laboratory, Ecosystem Center, Woods Hole MA 02543 1 Abstract Increasing atmospheric CO2 concentration will result in the acidifying and CO2 fertilization of our oceans. Phytoplankton assemblages will be among the organisms affected by theses changes. We tested how coastal New England phytoplankton community photosynthetic productivity and community composition will be changed. This microcosm experiment manipulated the CO2 concentrations to be 370µatm, 925µatm, and 3700µatm over a 3 week incubation period. Treatments showed reduced pH and changed dissolved inorganic carbon levels relative to each treatment. Measured parameters included nutrients, Chl a, pCO2, enumeration, and quantification of phytoplankton taxa using microscopy and flow cytometry. The present study revealed a coastal plankton community that was slightly affected by ocean acidification. Changes in respiration across the course of a 24 hour period showed net consumption cycles to be greatest at elevated CO2 levels. A species shift was seen from control treatments to elevated treatments and over a 5 day period. We have concluded that future research needs to be done to accurately determine how phytoplankton will be affected by ocean acidification changes. Key Words: Ocean acidification, phytoplankton, primary productivity, community composition Introduction Elevated atmospheric CO2 levels, primarily due to fossil fuel combustion, has led to increased CO2 fertilization of oceans (Brewer 2009), also known as ‘ocean acidification’. Currently the atmospheric CO2 concentration is 390ppm, and is expected to increase to 750 ppm or higher by the end of the century (Raven et al. 2005). Global elemental cycles are driven by biological activity. Assessing the impact of ocean acidification on marine microorganisms is important for understanding how aquatic systems will react. Observing microbe-driven ecosystem function changes to elevated CO2 has proven challenging (Liu et al. 2010) and (Joint et al. 2011). Phytoplankton play an important role as primary producers of aquatic systems. Direct effects between ocean acidification and photosynthetic ability have been observed to increase under elevated pCO2 (Rost et al. 2008). The net rate of organic carbon production determines support for higher trophic levels. Natural phytoplankton assemblages have been shown to enhance photosynthesis under elevated pCO2 (Egge et al. 2009). This study will focus on coastal phytoplankton community because they contribute significantly to global primary productivity (Field et al. 1998). Changes in community composition can also be caused by elevated pCO2. Global changes in biodiversity can alter ecosystem services and disturb biogeochemical cycles, such as control the CO2 taken up by the oceans. Different species play different roles in ecosystem 2 dynamics. For example, the size of phytoplankton determines grazing efficiencies and can alter the population structure of grazers. Microbial community structure shifts can mean a loss of biodiversity, compromised of ecosystem robustness, and potentially major consequences for higher trophic levels. Studies that have observed the effects of increased CO2 inputs on phytoplankton systems have found mixed results on phytoplankton assemblages (Nielsen et al. 2010). It is likely that different phytoplankton taxa react differently to ocean acidification. The effects of elevated pCO2 on microzooplankton have shown no consistent effect to ocean acidification on microbial biodiversity and community composition (Suffrian et al. 2008). In the following we describe the responses of natural fall coastal phytoplankton groups to changes in the carbonate chemistry of a microcosm system to determine how future lowered pH levels and increased CO2 may alter functioning and composition. This study focuses on the changes to phytoplankton photosynthetic productivity and community assemblages. To examine the effects we created semi-continuous microcosm culture techniques in which CO2 manipulated environments were periodically pulsed by pulling sampling and returning medium with nutrients and filtered seawater. Here, we report on the results of a 3 week experiment demonstrating effects of elevated CO2 levels of phytoplankton productivity on local North-west Atlantic Ocean assemblages. We discuss the potential ecological and biogeochemical implications of our findings. Methods A coastal seawater sample was obtained from Woods Hole, MA (41.5264° N, 70.6736° W) during November. The sample was filtered through 200µm mesh. 1L of sample was allocated to each microcosm with nutrients. Nutrient concentrations in the microcosms remained at 36µM KNO3, 52µM NaSiO3, and 2.3µM KH2PO4. The experiment was preformed on triplicates at 925µatm, 3700µatm, and controlled atmospheric levels of 370µatm. Microcosms were incubated at 20˚C on a 12 hour light cycle and were continuously stirred. CO2 and air input were bubbled into the sample at 23 mL min-1. Each day, 10% of the sample was removed and the same volume of 0.45µm filtered seawater and nutrients were added back in. All air and water samples were taken +/- 1 hour of growth lights turning on each day. The microcosms sat undisturbed for 4 days prior to initiation of the pulse chemostat method. 3 10mL of sample was used to measure pH with Accumet pH/conductivity meter (Fisher Scientific). 10mL was used for fluorescence using a Fluorometer (Turner Designs). 80mL of was filtered using 25mm GF/F filters for nutrients. Ammonium was measured using a modification of the phenol-hypochlorite method (Solorzano 1969) analyzed with a Cary UV Visible Spectrophotometer (Varian). Phosphate was analyzed by a modification of the method of Murphy and Riley (1962) using UV-VIS Spectrophotometer (Shimadzu). Nitrate was measured using QuickChem Flow Injection Analyzer (LACHET). Filters were dried, and analyzed for molar carbon and nitrogen with a PerkinElmer 2400 Series II CHN Elemental Analyzer. 10mL of sample was used to measure dissolved inorganic carbon levels. 20mL of Ascarite scrubbed CO2-free air was drawn into the syringe, 0.2mL of H2SO4 was added, and sample was shaken for 1 minute prior to injection of air. pCO2 in the microcosm head space and CO2 input to the system was also measured. CO2 consumption was calculated by subtracting -1 pCO2 input from output, then multiplying by flow rate (23mL min ). Dissolved inorganic carbon (DIC) and microcosm air was measured using gas chromatography (GC-8A Shimadzu). 50mL of sample was drawn for microscopy. Autotrophs were fixed in alkaline Lugol’s solution (10g iodine, 20g potassium iodide, 10g sodium acetate, in 140ml distilled water) for a concentration of 0.1%, followed by borate-buffered formalin addition of 2.4%, and 3% sodium thiosulfate for a final concentration of 0.1% (Sherr and Sherr, 1993). Preserved sample were filtered onto 25mm white 0.8µm membrane filters (Osmonics). Taxa were identified to the nearest genus or species. Identification was done using differential interference and bright field light microscopy (Zeiss Axio Imager.M2) at 20X and 40X. Additional samples (50mL) were fixed to a final concentration of 5% glutaric dialdehyde. Direct DAPI (4',6-diamino-2-phenylindole dihydrochloride) counts were taken from the glutaraldehyde preserved samples. 1 ml of sample and 50µl of 200µg/ml working solution DAPI was incubated for 5 minutes then drawn onto 1µm black polycarbonate filters. Phosphate buffered saline was used to rinse. Samples were viewed under 20X magnification using blue fluorescence. Samples were quantified: (cells/field of view)´(area of filter covered by sample) cells ml-1 = (field of view area)´(preseravtion dilution factor)´(ml filtered) Flow cytometry was done on live, unpreserved samples using a flow cytometer (BD FACSCalibur) using CellQuest Pro software. 1mL samples were drawn and filtered using 35µm 4 mesh, and 5µL of 1µm beads were added to the sample. Particles were enumerated based on size, complexity, and fluorescence. Results Measured pH during the experimental period remained constant once a steady pH was reached (Figure 1). There were significant differences in pH between treatments. The 370µatm (control) maintained a pH of 8.1, 925µatm was 7.6, and 3700µatm was 7.3. Seawater sampled at the start of the experiment measured a pH of 8. The initial level of DIC of the sampled water was 2700µM, and for all treatment levels the DIC changes occurred during the experimental period (Figure 2). Nutrient profiles phosphate and ammonium were measured throughout the course of the experiment. Initial concentrations of ammonium were undetectable, phosphate measured 0.66µM, and nitrate was also undetectable in the seawater sampled. Ammonium levels gradually increased in all treatments over the course of the experiment (Figure 3A). Phosphate levels gradually decreased in all treatments (Figure 3B). Levels of nitrate after day 5 were below detection limit (Figure 3C). No statistically significant differences were found between treatments. Chlorophyll a changes between treatments reflect changes based on treatment. Final Chl a measurements reflect significant differences between treatments, with the 3700µatm treatment having the highest Chl a values, and the ambient CO2 treatment measuring the lowest (Figure 4). Molar carbon and nitrogen levels were not significant between treatments, all treatments showed
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