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The responses of macroalgae to ocean acidification could be altered by availability of macronutrients, such as ammonium (NH4+). This study determined how the opportunistic macroalga, Ulva australis responded to simultaneous changes in decreasing pH and NH4+ enrichment. This was investigated in a week-long growth experiment across a range of predicted future pHs with ambient and enriched NH4+ treatments followed by measurements of relative growth rates (RGR), NH4+ 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 (rETRmax) and maximum quantum yield of photosystem II (PSII) photochemistry (Fv/Fm). Photosynthetic capacity was derived from the RLCs and included the efficiency of light harvesting (α), slope of photoinhibition (β), and the light saturation point (Ek). The results showed that NH4+ enrichment did not modify the effects of pH on RGRs, NH4+ uptake rates and pools, total chlorophyll, rETRmax, α, β, Fv/Fm, tissue C and N, and the C:N ratio. However, Ek was differentially affected by pH under different NH4+ treatments. Ek increased with decreasing pH in the ambient NH4+ treatment, but not in the enriched NH4+ treatment. NH4+ enrichment increased RGRs, NH4+ pools, total chlorophyll, rETRmax, α, β, Fv/Fm, and tissue N, and decreased NH4+ uptake rates and the C:N ratio. Decreased pH increased total chlorophyll content, rETRmax, Fv/Fm, and tissue N content, and decreased the C:N ratio. Therefore, the results indicate that U. australis growth is increased with NH4+ 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.

IntroductionThe coastal macroalgal genus, Ulva, is found worldwide and is considered a nuisance algal genus due to its propensity for forming vast blooms. The response of Ulva to ocean acidification (OA) is of concern, particularly with nutrient enrichment, as these combined drivers may enhance algal blooms because of increased availability of dissolved inorganic resources.MethodsWe determined how a suite of physiological parameters were affected by OA and ammonium (NH4+) enrichment in 22-day laboratory experiments to gain a mechanistic understanding of growth, nutrient assimilation, and photosynthetic processes. We predicted how physiological parameters change across a range of pCO2 and NH4+ scenarios to ascertain bloom potential under future climate change regimes.ResultsDuring the first five days of growth, there was a positive synergy between pCO2 and NH4+ enrichment, which could accelerate initiation of an Ulva bloom. After day 5, growth rates declined overall and there was no effect of pCO2, NH4+, nor their interaction. pCO2 and NH4+ acted synergistically to increase NO3- uptake rates, which may have contributed to increased growth in the first five days. Under the saturating photosynthetically active radiation (PAR) used in this experiment (500 μmol photon m-2 s-1), maximum photosynthetic rates were negatively affected by increased pCO2, which could be due to increased sensitivity to light when high CO2 reduces energy requirements for inorganic carbon acquisition. Activity of CCMs decreased under high pCO2 and high NH4+ conditions indicating that nutrients play a role in alleviating photodamage and regulating CCMs under high-light intensities.DiscussionThis study demonstrates that OA could play a role in initiating or enhancing Ulva blooms in a eutrophic environment and highlights the need for understanding the potential interactions among light, OA, and nutrient enrichment in regulating photosynthetic processes.