The complete mitochondrial genome of a wild-collected Kappaphycus malesianus (Solieriaceae, Rhodophyta).

Kappaphycus malesianus is a red seaweed farmed primarily for its carrageenan, a polysaccharide important in the food and pharmaceutical industries. Among the commercially cultivated Kappaphycus species, only K. malesianus has no mitogenome data available. Here, we assembled the mitochondrial genome of K. malesianus from next-generation sequencing data. The circular mitogenome consisted of 25,250 base pairs (bp) with a GC content of 30.25%. These values were comparable to previously sequenced solieriacean mitogenomes. Structural features, such as the stem-loop and hairpin, which were previously reported in other rhodophytes mitochondrial DNA, were also identified. The annotated genes (24 protein-coding genes, 24 tRNA genes, and 2 rRNA genes) were arranged in an order similar to the other available solieriacean mitogenomes. Lastly, phylogenetic analysis using 23 predicted protein domains showed the sister relationship of K. malesianus with other Kappaphycus species.

Growth, nitrate uptake kinetics, and biofiltration potential of eucheumatoids with different thallus morphologies.

The declining production of commercially important eucheumatoids related to serious problems, like increasing susceptibility to ice-ice disease and epiphytism, may be ameliorated by nutrition. This ushered an increasing interest in incorporating seaweeds into an integrated multi-trophic aquaculture (IMTA) setup to take up excess inorganic nutrients produced by fish farms for their nourishment. In this regard, it is important to understand the nutrient uptake capacity of candidate seaweeds for incorporation into an IMTA system. Here, we examined the growth, nitrate ( NO3- ) uptake kinetics, and biofiltration potential of Eucheuma denticulatum and three strains of Kappaphycus alvarezii (G-O2, TR-C16, and SW-13) with distinct thallus morphologies. The NO3- uptake rates of the samples were determined under a range of NO3- concentrations (1-48 µM) and uptake rates were fitted to the Michaelis-Menten saturation equation. Among the examined eucheumatoids, only SW-13 had a linear response to NO3- concentration while other strains had uptake rates that followed the Michaelis-Menten saturation equation. Eucheuma denticulatum had the lowest Km (9.78 ± 1.48 µM) while G-O2 had the highest Vmax (307 ± 79.3 µmol · g-1 · min-1 ). The efficiency in NO3- uptake (highest Vmax /Km and α) was translated into the highest growth rate (3.41 ± 0.58% · d-1 ) measured in E. denticulatum. Our study provided evidence that eucheumatoids could potentially take up large amount of NO3- and fix CO2 when cultivated proximate to a fish farm as one component of an IMTA system. During a 45 -d cultivation period of eucheumatoids, as much as 370 g NO3- can be sequestered by every 1 kg initial biomass of E. denticulatum growing at 3% · d-1 . Furthermore, based on our unpublished photosynthetic measurements, the congeneric K. striatus can fix 27.5 g C · kg-1 DW during a 12 h daylight period.

Reproductive phenology and morphology of Macrocystis pyrifera (Laminariales, Ochrophyta) from southern New Zealand in relation to wave exposure1.

Macrocystis pyrifera is a major habitat forming kelp in coastal ecosystems of temperate regions of the northern and southern hemispheres. We investigated the seasonal occurrence of adult sporophytes, morphological characteristics, and reproductive phenology at two sites within a wave-protected harbour and two wave-exposed sites in southern New Zealand every 3-4 months between 2012 and 2013. Seasonality in reproduction was assessed via the number of sporophylls, the occurrence of sori on sporophylls, and non-sporophyllous laminae (fertile pneumatocyst-bearing blades and fertile apical scimitars), meiospore release, and germination. We found that M. pyrifera was present and reproductive year-round in three of the four sites, and patterns were similar for the wave-exposure conditions. Sori were found on pneumatocyst-bearing blades and apical scimitars in addition to the sporophylls, and viable meiospores were released from all three types of laminae. Morphological variations between sites with different wave exposure indicate that sporophytes from wave-protected sites have bigger blades and holdfasts and are longer than those from wave-exposed sites. We discuss the implications of these biological variables for the ecology of M. pyrifera inhabiting different wave exposure environments in southern New Zealand.

Chemical profiling of Ulva species for food applications: What is in a name?

Seaweeds are increasingly used in European cuisine. Until the recent use of molecular techniques, species identification was solely based on morphology which cannot easily discriminate morphologically simple but phenotypically plastic taxa such as the green algal genus Ulva. For example, current taxonomic protocol effectively reassigned the previously known European 'Ulva lactuca L.' under the name Ulva fenestrata Postels & Ruprecht. Also, the presumptive Ulva lactuca approved by the Institute for Reference Materials and Measurements (IRMM, Joint Research Center, European Commission) as Certified Reference Material (CRM) for analytical quality assurance was genetically identified as U. rigida C.Agardh. It is very likely that different Ulva species under various names have been consumed as food not only in Europe, but also worldwide. In this regard, when chemical composition and nutritional quality of different seaweed species meet a set of food standard criteria, and food safety hazards are mitigated, they should be endorsed for consumption. In the case of Ulva, we propose that different bladed and tubular species should generally be accepted for food consumption in Europe.

Chemical profiling of the Arctic sea lettuce Ulva lactuca (Chlorophyta) mass-cultivated on land under controlled conditions for food applications.

The increasing use of seaweeds in European cuisine led to cultivation initiatives funded by the European Union. Ulva lactuca, commonly known as sea lettuce, is a fast growing seaweed in the North Atlantic that chefs are bringing into the local cuisine. Here, different strains of Arctic U. lactuca were mass-cultivated under controlled conditions for up to 10 months. We quantified various chemical constituents associated with both health benefits (carbohydrates, protein, fatty acids, minerals) and health risks (heavy metals). Chemical analyses showed that long-term cultivation provided biomass of consistently high food quality and nutritional value. Concentrations of macroelements (C, N, P, Ca, Na, K, Mg) and micronutrients (Fe, Zn, Co, Mn, I) were sufficient to contribute to daily dietary mineral intake. Heavy metals (As, Cd, Hg and Pb) were found at low levels to pose health risk. The nutritional value of Ulva in terms of carbohydrates, protein and fatty acids is comparable to some selected fruits, vegetables, nuts and grains.

Heat stress responses and population genetics of the kelp Laminaria digitata (Phaeophyceae) across latitudes reveal differentiation among North Atlantic populations.

To understand the thermal plasticity of a coastal foundation species across its latitudinal distribution, we assess physiological responses to high temperature stress in the kelp Laminaria digitata in combination with population genetic characteristics and relate heat resilience to genetic features and phylogeography. We hypothesize that populations from Arctic and cold-temperate locations are less heat resilient than populations from warm distributional edges. Using meristems of natural L. digitata populations from six locations ranging between Kongsfjorden, Spitsbergen (79°N), and Quiberon, France (47°N), we performed a common-garden heat stress experiment applying 15°C to 23°C over eight days. We assessed growth, photosynthetic quantum yield, carbon and nitrogen storage, and xanthophyll pigment contents as response traits. Population connectivity and genetic diversity were analyzed with microsatellite markers. Results from the heat stress experiment suggest that the upper temperature limit of L. digitata is nearly identical across its distribution range, but subtle differences in growth and stress responses were revealed for three populations from the species' ecological range margins. Two populations at the species' warm distribution limit showed higher temperature tolerance compared to other populations in growth at 19°C and recovery from 21°C (Quiberon, France), and photosynthetic quantum yield and xanthophyll pigment responses at 23°C (Helgoland, Germany). In L. digitata from the northernmost population (Spitsbergen, Norway), quantum yield indicated the highest heat sensitivity. Microsatellite genotyping revealed all sampled populations to be genetically distinct, with a strong hierarchical structure between southern and northern clades. Genetic diversity was lowest in the isolated population of the North Sea island of Helgoland and highest in Roscoff in the English Channel. All together, these results support the hypothesis of moderate local differentiation across L. digitata's European distribution, whereas effects are likely too weak to ameliorate the species' capacity to withstand ocean warming and marine heatwaves at the southern range edge.

Inorganic carbon utilization of tropical calcifying macroalgae and the impacts of intensive mariculture-derived coastal acidification on the physiological performance of the rhodolith Sporolithon sp.

Fish farming in coastal areas has become an important source of food to support the world's increasing population. However, intensive and unregulated mariculture activities have contributed to changing seawater carbonate chemistry through the production of high levels of respiratory CO2. This additional CO2, i.e. in addition to atmospheric inputs, intensifies the effects of global ocean acidification resulting in localized extreme low pH levels. Marine calcifying macroalgae are susceptible to such changes due to their CaCO3 skeleton. Their physiological response to CO2-driven acidification is dependent on their carbon physiology. In this study, we used the pH drift experiment to determine the capability of 9 calcifying macroalgae to use one or more inorganic carbon (Ci) species. From the 9 species, we selected the rhodolith Sporolithon sp. as a model organism to investigate the long-term effects of extreme low pH on the physiology and biochemistry of calcifying macroalgae. Samples were incubated under two pH treatments (pH 7.9 = ambient and pH 7.5 = extreme acidification) in a temperature-controlled (26 ± 0.02 °C) room provided with saturating light intensity (98.3 ± 2.50 µmol photons m-2 s-1). After the experimental treatment period (40 d), growth rate, calcification rate, nutrient uptake rate, organic content, skeletal CO3-2, pigments, and tissue C, N and P of Sporolithon samples were compared. The pH drift experiment revealed species-specific Ci use mechanisms, even between congenerics, among tropical calcifying macroalgae. Furthermore, long-term extreme low pH significantly reduced the growth rate, calcification rate and skeletal CO3-2 content by 79%, 66% and 18%, respectively. On the other hand, nutrient uptake rates, organic matter, pigments and tissue C, N and P were not affected by the low pH treatments. Our results suggest that the rhodolith Sporolithon sp. is susceptible to the negative effects of extreme low pH resulting from intensive mariculture-driven coastal acidification.

Effects of multiple drivers of ocean global change on the physiology and functional gene expression of the coccolithophore Emiliania huxleyi.

Ongoing ocean global change due to anthropogenic activities is causing multiple chemical and physical seawater properties to change simultaneously, which may affect the physiology of marine phytoplankton. The coccolithophore Emiliania huxleyi is a model species often employed in the study of the marine carbon cycle. The effect of ocean acidification (OA) on coccolithophore calcification has been extensively studied; however, physiological responses to multiple environmental drivers are still largely unknown. Here we examined two-way and multiple driver effects of OA and other key environmental drivers-nitrate, phosphate, irradiance, and temperature-on the growth, photosynthetic, and calcification rates, and the elemental composition of E. huxleyi. In addition, changes in functional gene expression were examined to understand the molecular mechanisms underpinning the physiological responses. The single driver manipulation experiments suggest decreased nitrate supply being the most important driver regulating E. huxleyi physiology, by significantly reducing the growth, photosynthetic, and calcification rates. In addition, the interaction of OA and decreased nitrate supply (projected for year 2100) had more negative synergistic effects on E. huxleyi physiology than all other two-way factorial manipulations, suggesting a linkage between the single dominant driver (nitrate) effects and interactive effects with other drivers. Simultaneous manipulation of all five environmental drivers to the conditions of the projected year 2100 had the largest negative effects on most of the physiological metrics. Furthermore, functional genes associated with inorganic carbon acquisition (RubisCO, AEL1, and δCA) and calcification (CAX3, AEL1, PATP, and NhaA2) were most downregulated by the multiple driver manipulation, revealing linkages between responses of functional gene expression and associated physiological metrics. These findings together indicate that for more holistic projections of coccolithophore responses to future ocean global change, it is necessary to understand the relative importance of environmental drivers both individually (i.e., mechanistic understanding) and interactively (i.e., cumulative effect) on coccolithophore physiology.

Variability and Potential of Seaweeds as Ingredients of Ruminant Diets: An In Vitro Study.

This study was designed to analyze the chemical composition and in vitro rumen fermentation of eight seaweed species (Brown: Alaria esculenta, Laminaria digitata, Pelvetia canaliculata, Saccharina latissima; Red: Mastocarpus stellatus, Palmaria palmata and Porphyra sp.; Green: Cladophora rupestris) collected in Norway during spring and autumn. Moreover, the in vitro ruminal fermentation of seventeen diets composed of 1:1 oat hay:concentrate, without (control diet) or including seaweeds was studied. The ash and N contents were greater (p < 0.001) in seaweeds collected during spring than in autumn, but autumn-seaweeds had greater total extractable polyphenols. Nitrogen in red and green seaweeds was greater than 2.20 and in brown seaweeds, it was lower than 1.92 g/kg DM. Degradability after 24 h of fermentation was greater in spring seaweeds than in autumn, with Palmaria palmata showing the greatest value and Pelvetia canaliculata the lowest. Seaweeds differed in their fermentation pattern, and autumn Alaria esculenta, Laminaria digitata, Saccharina latissima and Palmaria palmata were similar to high-starch feeds. The inclusion of seaweeds in the concentrate of a diet up to 200 g/kg concentrate produced only subtle effects on in vitro ruminal fermentation.

Tolerance of Fucus vesiculosus exposed to diesel water-accommodated fraction (WAF) and degradation of hydrocarbons by the associated bacteria.

The viability and physiological state of brown macroalgae Fucus vesiculosus and its associated epiphytic bacteria exposed to diesel water-accommodated fraction (WAF), as well as the capacity of this association to deplete petroleum hydrocarbons (HCs) were experimentally tested. After a 6-day exposure treatment, the algal-surface associated bacteria were identified as primarily hydrocarbon-oxidising bacteria (HOB), and the algal-HOB association was able to deplete petroleum hydrocarbons from the diesel WAF by 80%. The HOB density on the algal surface exposed to diesel WAF was 350% higher compared to the control (i.e. HOB density on the algal surface exposed to ambient seawater), which suggest that they actively proliferated in the presence of hydrocarbons and most likely consumed hydrocarbons as their primary organic substrate. Exposure to diesel WAF did not affect the metabolic activity of F. vesiculosus. Higher lipid peroxidation was observed in F. vesiculosus exposed to diesel WAF while catalase concentration decreased only during the first day of exposure. Results suggest F. vesiculosus is tolerant to oil pollution and the algal-HOB association can efficiently deplete petroleum hydrocarbons in oil-contaminated seas.

Copper pollution exacerbates the effects of ocean acidification and warming on kelp microscopic early life stages.

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.

Iodine content in bulk biomass of wild-harvested and cultivated edible seaweeds: Inherent variations determine species-specific daily allowable consumption.

This study represents a large-scale investigation into iodine contents in three commercially important and edible seaweed species from the North Atlantic: the brown algae Saccharina latissima and Alaria esculenta, and the red alga Palmaria palmata. Variability among and within species were explored in terms of temporal and spatial variations in addition to biomass source. Mean iodine concentration in bulk seaweed biomass was species-specific: Saccharina > Alaria > Palmaria. Iodine contents of Saccharina biomass were similar between years and seasons, but varied significantly between sampling locations and biomass sources. In Alaria and Palmaria, none of the independent variables examined contributed significantly to the small variations observed. Our data suggest that all three species are rich sources of iodine, and only 32, 283, or 2149 mg dry weight of unprocessed dry biomass of Saccharina, Alaria, or Palmaria, respectively, meets the recommended daily intake levels for most healthy humans.

Ocean acidification and kelp development: Reduced pH has no negative effects on meiospore germination and gametophyte development of Macrocystis pyrifera and Undaria pinnatifida.

The absorption of anthropogenic CO2 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 (pHT 7.20, extreme OA predicted for 2300; pHT 7.65, OA predicted for 2100; pHT 8.01, ambient pH; and pHT 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 pHT (7.20 and 7.65) had positive effects on germling growth rate and gametophyte size in both M. pyrifera and U. pinnatifida, whereas, higher pHT (8.01 and 8.40) reduced the gametophyte size in both kelps. Sex ratio of gametophytes of both kelps was biased toward females under all pHT treatments, except for U. pinnatifida at pHT 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 pHT treatments, indicating that the microscopic stages of the native M. pyrifera and the invasive U. pinnatifida will respond similarly to OA.

Seawater pH, and not inorganic nitrogen source, affects pH at the blade surface of Macrocystis pyrifera: implications for responses of the giant kelp to future oceanic conditions.

Ocean acidification (OA), the ongoing decline in seawater pH, is predicted to have wide-ranging effects on marine organisms and ecosystems. For seaweeds, the pH at the thallus surface, within the diffusion boundary layer (DBL), is one of the factors controlling their response to OA. Surface pH is controlled by both the pH of the bulk seawater and by the seaweeds' metabolism: photosynthesis and respiration increase and decrease pH within the DBL (pHDBL ), respectively. However, other metabolic processes, especially the uptake of inorganic nitrogen (Ni ; NO3- and NH4+ ) may also affect the pHDBL . Using Macrocystis pyrifera, we hypothesized that (1) NO3- uptake will increase the pHDBL , whereas NH4+ uptake will decrease it, (2) if NO3- is cotransported with H+ , increases in pHDBL would be greater under an OA treatment (pH = 7.65) than under an ambient treatment (pH = 8.00), and (3) decreases in pHDBL will be smaller at pH 7.65 than at pH 8.00, as higher external [H+ ] might affect the strength of the diffusion gradient. Overall, Ni source did not affect the pHDBL . However, increases in pHDBL were greater at pH 7.65 than at pH 8.00. CO2 uptake was higher at pH 7.65 than at pH 8.00, whereas HCO3- uptake was unaffected by pH. Photosynthesis and respiration control pHDBL rather than Ni uptake. We suggest that under future OA, Macrocystis pyrifera will metabolically modify its surface microenvironment such that the physiological processes of photosynthesis and Ni uptake will not be affected by a reduced pH.

Effect of Ocean Acidification and pH Fluctuations on the Growth and Development of Coralline Algal Recruits, and an Associated Benthic Algal Assemblage.

Coralline algae are susceptible to the changes in the seawater carbonate system associated with ocean acidification (OA). However, the coastal environments in which corallines grow are subject to large daily pH fluctuations which may affect their responses to OA. Here, we followed the growth and development of the juvenile coralline alga Arthrocardia corymbosa, which had recruited into experimental conditions during a prior experiment, using a novel OA laboratory culture system to simulate the pH fluctuations observed within a kelp forest. Microscopic life history stages are considered more susceptible to environmental stress than adult stages; we compared the responses of newly recruited A. corymbosa to static and fluctuating seawater pH with those of their field-collected parents. Recruits were cultivated for 16 weeks under static pH 8.05 and 7.65, representing ambient and 4× preindustrial pCO2 concentrations, respectively, and two fluctuating pH treatments of daily [Formula: see text] (daytime pH = 8.45, night-time pH = 7.65) and daily [Formula: see text] (daytime pH = 8.05, night-time pH = 7.25). Positive growth rates of new recruits were recorded in all treatments, and were highest under static pH 8.05 and lowest under fluctuating pH 7.65. This pattern was similar to the adults' response, except that adults had zero growth under fluctuating pH 7.65. The % dry weight of MgCO3 in calcite of the juveniles was reduced from 10% at pH 8.05 to 8% at pH 7.65, but there was no effect of pH fluctuation. A wide range of fleshy macroalgae and at least 6 species of benthic diatoms recruited across all experimental treatments, from cryptic spores associated with the adult A. corymbosa. There was no effect of experimental treatment on the growth of the benthic diatoms. On the community level, pH-sensitive species may survive lower pH in the presence of diatoms and fleshy macroalgae, whose high metabolic activity may raise the pH of the local microhabitat.

Long-Term Conditioning to Elevated pCO2 and Warming Influences the Fatty and Amino Acid Composition of the Diatom Cylindrotheca fusiformis.

The unabated rise in anthropogenic CO2 emissions is predicted to strongly influence the ocean's environment, increasing the mean sea-surface temperature by 4°C and causing a pH decline of 0.3 units by the year 2100. These changes are likely to affect the nutritional value of marine food sources since temperature and CO2 can influence the fatty (FA) and amino acid (AA) composition of marine primary producers. Here, essential amino (EA) and polyunsaturated fatty (PUFA) acids are of particular importance due to their nutritional value to higher trophic levels. In order to determine the interactive effects of CO2 and temperature on the nutritional quality of a primary producer, we analyzed the relative PUFA and EA composition of the diatom Cylindrotheca fusiformis cultured under a factorial matrix of 2 temperatures (14 and 19°C) and 3 partial pressures of CO2 (180, 380, 750 µatm) for >250 generations. Our results show a decay of ~3% and ~6% in PUFA and EA content in algae kept at a pCO2 of 750 µatm (high) compared to the 380 µatm (intermediate) CO2 treatments at 14°C. Cultures kept at 19°C displayed a ~3% lower PUFA content under high compared to intermediate pCO2, while EA did not show differences between treatments. Algae grown at a pCO2 of 180 µatm (low) had a lower PUFA and AA content in relation to those at intermediate and high CO2 levels at 14°C, but there were no differences in EA at 19°C for any CO2 treatment. This study is the first to report adverse effects of warming and acidification on the EA of a primary producer, and corroborates previous observations of negative effects of these stressors on PUFA. Considering that only ~20% of essential biomolecules such as PUFA (and possibly EA) are incorporated into new biomass at the next trophic level, the potential impacts of adverse effects of ocean warming and acidification at the base of the food web may be amplified towards higher trophic levels, which rely on them as source of essential biomolecules.

Effects of ocean acidification on the photosynthetic performance, carbonic anhydrase activity and growth of the giant kelp Macrocystis pyrifera.

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.

Saturating light and not increased carbon dioxide under ocean acidification drives photosynthesis and growth in Ulva rigida (Chlorophyta).

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 [Formula: see text] dehydration and alter the stable carbon isotope (δ (13)C) 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 [Formula: see text] use mechanisms, that is, direct [Formula: see text] uptake through the AE port and CAext-mediated [Formula: see text] 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 [Formula: see text] transporters to which the remaining NPS can be attributed. The shift in δ (13)C signatures from -22‰ toward -10‰ under saturating light but not under elevated CO2(aq) suggest preference and substantial [Formula: see text] 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.