Under ocean acidification (OA), the 200 % increase in CO2(aq) and the reduction of pH by 0.3-0.4 units are predicted to affect the carbon physiology and growth of macroalgae. Here we examined how the physiology of the giant kelp Macrocystis pyrifera is affected by elevated pCO2/low pH. Growth and photosynthetic rates, external and internal carbonic anhydrase (CA) activity, HCO3 (-) versus CO2 use were determined over a 7-day incubation at ambient pCO2 400 µatm/pH 8.00 and a future OA treatment of pCO2 1200 µatm/pH 7.59. Neither the photosynthetic nor growth rates were changed by elevated CO2 supply in the OA treatment. These results were explained by the greater use of HCO3 (-) compared to CO2 as an inorganic carbon (Ci) source to support photosynthesis. Macrocystis is a mixed HCO3 (-) and CO2 user that exhibits two effective mechanisms for HCO3 (-) utilization; as predicted for species that possess carbon-concentrating mechanisms (CCMs), photosynthesis was not substantially affected by elevated pCO2. The internal CA activity was also unaffected by OA, and it remained high and active throughout the experiment; this suggests that HCO3 (-) uptake via an anion exchange protein was not affected by OA. Our results suggest that photosynthetic Ci uptake and growth of Macrocystis will not be affected by elevated pCO2/low pH predicted for the future, but the combined effects with other environmental factors like temperature and nutrient availability could change the physiological response of Macrocystis to OA. Therefore, further studies will be important to elucidate how this species might respond to the global environmental change predicted for the ocean.

Carbon physiology of a genetically identified Ulva rigida was investigated under different CO2(aq) and light levels. The study was designed to answer whether (1) light or exogenous inorganic carbon (Ci) pool is driving growth; and (2) elevated CO2(aq) concentration under ocean acidification (OA) will downregulate CAext-mediated dehydration and alter the stable carbon isotope (δ13C) signatures toward more CO2 use to support higher growth rate. At pHT 9.0 where CO2(aq) is <1 μmol L−1, inhibition of the known use mechanisms, that is, direct uptake through the AE port and CAext-mediated dehydration decreased net photosynthesis (NPS) by only 56–83%, leaving the carbon uptake mechanism for the remaining 17–44% of the NPS unaccounted. An in silico search for carbon-concentrating mechanism elements in expressed sequence tag libraries of Ulva found putative light-dependent transporters to which the remaining NPS can be attributed. The shift in δ13C signatures from –22‰ toward –10‰ under saturating light but not under elevated CO2(aq) suggest preference and substantial use to support photosynthesis and growth. U. rigida is Ci saturated, and growth was primarily controlled by light. Therefore, increased levels of CO2(aq) predicted for the future will not, in isolation, stimulate Ulva blooms.

Macrocystis pyrifera is a widely distributed, highly productive, seaweed. It is known to use bicarbonate (HCO3 (-) ) from seawater in photosynthesis and the main mechanism of utilization is attributed to the external catalyzed dehydration of HCO3 (-) by the surface-bound enzyme carbonic anhydrase (CAext ). Here, we examined other putative HCO3 (-) uptake mechanisms in M. pyrifera under pHT 9.00 (HCO3 (-) : CO2  = 940:1) and pHT 7.65 (HCO3 (-) : CO2  = 51:1). Rates of photosynthesis, and internal CA (CAint ) and CAext activity were measured following the application of AZ which inhibits CAext , and DIDS which inhibits a different HCO3 (-) uptake system, via an anion exchange (AE) protein. We found that the main mechanism of HCO3 (-) uptake by M. pyrifera is via an AE protein, regardless of the HCO3 (-) : CO2 ratio, with CAext making little contribution. Inhibiting the AE protein led to a 55%-65% decrease in photosynthetic rates. Inhibiting both the AE protein and CAext at pHT 9.00 led to 80%-100% inhibition of photosynthesis, whereas at pHT 7.65, passive CO2 diffusion supported 33% of photosynthesis. CAint was active at pHT 7.65 and 9.00, and activity was always higher than CAext , because of its role in dehydrating HCO3 (-) to supply CO2 to RuBisCO. Interestingly, the main mechanism of HCO3 (-) uptake in M. pyrifera was different than that in other Laminariales studied (CAext -catalyzed reaction) and we suggest that species-specific knowledge of carbon uptake mechanisms is required in order to elucidate how seaweeds might respond to future changes in HCO3 (-) :CO2 due to ocean acidification.

Local and global changes associated with anthropogenic activities are impacting marine and terrestrial ecosystems. Macroalgae, especially habitat-forming species like kelp, play critical roles in temperate coastal ecosystems. However, their abundance and distribution patterns have been negatively affected by warming in many regions around the globe. Along with global change, coastal ecosystems are also impacted by local drivers such as eutrophication. The interaction between global and local drivers might modulate kelp responses to environmental change. This study examines the regulatory effect of no 3 − on the thermal plasticity of the giant kelp Macrocystis pyrifera. To do this, thermal performance curves (TPCs) of key temperature-dependant traits-growth, photosynthesis, NO 3 − assimilation and chlorophyll a fluorescence-were examined under nitrate replete and deplete conditions in a short-term incubation. We found that thermal plasticity was modulated by NO 3 − but different thermal responses were observed among traits. Our study reveals that nitrogen, a local driver, modulates kelp responses to high seawater temperatures, ameliorating the negative impacts on physiological performance (i.e. growth and photosynthesis). However, this effect might be species-specific and vary among biogeographic regions -thus, further work is needed to determine the generality of our findings to other key temperate macroalgae that are experiencing temperatures close to their thermal tolerance due to climate change.Rising levels of atmospheric CO 2 are causing increases in air and sea surface temperatures (SSTs), with the mean SST predicted to rise by 1.4 °C to 4.8 °C by 2100 1 . With global warming, extreme high temperature events such as marine heat waves (MHWs) have also increased in frequency, intensity and duration along the World's coastline, including the Mediterranean, Australia and Brazilian Atlantic sea 2-6 . These anomalous elevated temperatures have negatively impacted marine and terrestrial ecosystems by altering species' composition and distribution patterns 7-9 . Such ecological changes are also severely impacting ecosystem goods and services such as fisheries, and carbon sequestration and storage 10 . The impacts of warming are considerably larger in marine systems because of their greater sensitivity to these global stressors compared to terrestrial systems 11,12 . Because of this, there is rising concern about the capacity of marine species to acclimate quickly enough to short-term variability in temperature, which will be critical for organisms to adapt and survive in a changing ocean 13,14 .In ectothermic organisms such as plants, algae, invertebrates and lower vertebrates, temperature is the major factor regulating their physiology, growth, performance and fitness [15][16][17][18][19][20] . Therefore, changes in environmental temperatures (T a ) due to climate change will trigger modifications at physiological and biochemical levels, influencing whole-organism thermal plasticity (i.e. thermal sensitivities and...

Ocean warming (OW), ocean acidification (OA) and their interaction with local drivers, e.g., copper pollution, may negatively affect macroalgae and their microscopic life stages. We evaluated meiospore development of the kelps Macrocystis pyrifera and Undaria pinnatifida exposed to a factorial combination of current and 2100-predicted temperature (12 and 16 °C, respectively), pH (8.16 and 7.65, respectively), and two copper levels (no-added-copper and species-specific germination Cu-EC50). Meiospore germination for both species declined by 5–18% under OA and ambient temperature/OA conditions, irrespective of copper exposure. Germling growth rate declined by >40%·day−1, and gametophyte development was inhibited under Cu-EC50 exposure, compared to the no-added-copper treatment, irrespective of pH and temperature. Following the removal of copper and 9-day recovery under respective pH and temperature treatments, germling growth rates increased by 8–18%·day−1. The exception was U. pinnatifida under OW/OA, where growth rate remained at 10%·day−1 before and after copper exposure. Copper-binding ligand concentrations were higher in copper-exposed cultures of both species, suggesting that ligands may act as a defence mechanism of kelp early life stages against copper toxicity. Our study demonstrated that copper pollution is more important than global climate drivers in controlling meiospore development in kelps as it disrupts the completion of their life cycle.

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Ocean acidification and kelp development: Reduced pH has no negative effects on meiospore germination and gametophyte development of Macrocystis pyrifera and Undaria pinnatifida Accepted Article Accepted ArticleThis article is protected by copyright. All rights reserved. ABSTRACTThe absorption of anthropogenic CO 2 by the oceans is causing a reduction in the pH of the surface waters termed ocean acidification (OA). This could have substantial effects on marine coastal environments where fleshy (non-calcareous) macroalgae are dominant primary producers and ecosystem engineers. Few OA studies have focused on the early life stages of large macroalgae such as kelps. This study evaluated the effects of seawater pH on the ontogenic development of meiospores of the native kelp Macrocystis pyrifera and the invasive kelp Undaria pinnatifida, in south-eastern New Zealand. Meiospores of both kelps were released into four seawater pH treatments (pH T 7.20, extreme OA predicted for 2300; pH T 7.65, OA predicted for 2100; pH T 8.01, ambient pH; and pH T 8.40, pre-industrial pH) and cultured for 15 d. Meiospore germination, germling growth rate, and gametophyte size and sex ratio were monitored and measured. Exposure to reduced pH T (7.20 and 7.65) had Accepted Article Accepted ArticleThis article is protected by copyright. All rights reserved.positive effects on germling growth rate and gametophyte size in both M. pyrifera and U. pinnatifida, whereas, higher pH T (8.01 and 8.40) reduced the gametophyte size in both kelps.Sex ratio of gametophytes of both kelps was biased towards females under all pH T treatments, except for U. pinnatifida at pH T 7.65. Germling growth rate under OA was significantly higher in M. pyrifera compared to U. pinnatifida but gametophyte development was equal for both kelps under all seawater pH T treatments, indicating that the microscopic stages of the native M. pyrifera and the invasive U. pinnatifida will respond similarly to OA.Keywords: gametophyte, germination, hydrogen ion concentration, Laminariales, microscopic stages, Macrocystis, Undaria Abbreviations: ANOVA, analyses of variances; CA, carbonic anhydrase; CCMs, carbon concentrating mechanisms; CO 2 , carbon dioxide; H + , hydrogen proton; HCl, hydrochloric acid; HCO 3 -, bicarbonate; NaHCO 3 , sodium bicarbonate; NaOH, sodium hydroxide; OA, ocean acidification; pH T , pH measured on the total scale; AT, total alkalinity; DIC, dissolved inorganic carbon INTRODUCTIONMarine fleshy (non-calcifying) macroalgae are important components of coastal environments due to their high productivity and ecological function as ecosystem engineers, a source of food to higher trophic levels and refuge providers (Dayton 1985, Graham et al. 2007, Hurd et al. 2014. Macroalgal cells fix CO 2 using RuBisCO but due to its low affinity for CO 2 (Raven 1997, Giordano et al. 2005, most macroalgae have developed carbon Accepted Article Accepted ArticleThis article is protected by copyright. All rights reserved.concentrating mechanisms (CCMs). In macroalgae with ...