Multi-scale modeling of intensive macroalgae cultivation and marine nitrogen sequestration.

Multi-scale macroalgae growth models are required for the efficient design of sustainable, economically viable, and environmentally safe farms. Here, we develop a multi-scale model for Ulva sp. macroalgae growth and nitrogen sequestration in an intensive cultivation farm, regulated by temperature, light, and nutrients. The model incorporates a range of scales by incorporating spatial effects in two steps: light extinction at the reactor scale (1 m) and nutrient absorption at the farm scale (1 km). The model was validated on real data from an experimental reactor installed in the sea. Biomass production rates, chemical compositions, and nitrogen removal were simulated under different seasons, levels of dilution in the environment and water-exchange rate in the reactor. This multi-scale model provides an important tool for environmental authorities and seaweed farmers who desire to upscale to large bioremediation and/or macroalgae biomass production farms, thus promoting the marine sustainable development and the macroalgae-based bioeconomy.

Synthesis and release of fatty acids under the interaction of Ulva pertusa and Heterosigma akashiwo by stable isotope analysis.

Symbiosis of marine algae is inevitable in the marine environment, and species may occur interaction on the growth. In this study, the macroalgae Ulva pertusa and marine microalgae Heterosigma akashiwo were selected as target species to study the interaction mechanism between them. After the 8 days of co-cultivation, the inhibition on growth was observed for both of U. pertusa and H. akashiwo. Eight fatty acids in U. pertusa was detected, with the significant decrease in contents of polyunsaturated fatty acids (PUFAs) especially for C18:2, C18:3n-3 and C18:3n-6. Twelve fatty acids in H. akashiwo was detected, with the significant change for PUFAs. PUFA concentrations in the co-culture group were less than those in the mono-culture. Meanwhile the principal component analysis was conducted to insight into the interaction between U. pertusa and H. akashiwo by fatty acids content and carbon stable isotope ratio of fatty acids (δ13CFAs). Fatty acid content could not distinguish mono and co-culture. However, δ13CFAs could distinguish not only the culture time of algae, but also the living environment of algae. In addition, this study combined fatty acids content and δ13CFAs to explore the release of fatty acids by algae into the seawater. The C18:3n-3 was identified as the allelochemical released by U. pertusa to inhibit the growth of H. akashiwo. The ratio of δ13CFAs in seawater decreased. This study provides a theoretical basis for the symbiosis of marine algae, and a new method of compound-specific stable carbon isotopes was used to better explore the metabolism of fatty acids in algae.

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.

X-ray Fluorescence Techniques in Determining the Habitat Preferences of Species-Ulva pilifera (Ulvales, Chlorophyta) from Montenegro Case Study.

The paper presents four new sites where bright green Ulva thalli were found inhabiting freshwater (a river, a ditch, the Milet Canal) and marine (on the rocky shore of the Adriatic Sea) habitats in Montenegro. The aims of this study were to determine, for the first time, whether specimens of Ulva pilifera collected in Montenegro are phylogenetically and morphologically the same species as the one occurring in Europe. Using total reflection X-ray fluorescence (TXRF) and wavelength dispersive X-ray fluorescence (WDXRF) techniques it assessed the elemental composition of their thalli and its influence to colonise new habitats. Elements: Al, As, Ba, Br, Ca, Cl, Cr, Cu, Fe, Hf, I, K, Mg, Mn, Na, Ni, P, Pb, Rb, S, Si, Sr, Ti, V, and Zn were determined. The highest elemental concentrations were found for Ca = 16.3% (using WDXRF) and for Sr = 292 ppm (using TXRF) in the Ulva thalli. Ulva pilifera analysed from Montenegro, based on classical morphological methods and molecular techniques, are closely related to the same species from inland and coastal waters throughout Europe. The analysis of trace elements showed that the metal content in Ulva thalli is correlated with the trace elements in water and sediments. Ulva pilifera fits numerous features that make it one of the bioindicators of marine pollution, thanks to its worldwide distribution and capacity to accumulate trace elements.

Fourfold daily growth rate in multicellular marine alga Ulva meridionalis.

Microalgae with high growth rates have been considered as promising organisms to replace fossil resources with contemporary primary production as a renewable source. However, their microscopic size makes it hard to be harvested for industrial applications. In this regard, multicellular macroalgae are more suitable for harvesting. Here, we show that Ulva meridionalis has the highest growth rate ever reported for a multicellular autotrophic plant. Contrasted to the known bloom-forming species U. prolifera growing at an approximately two-fold growth rate per day in optimum conditions, U. meridionalis grows at a daily rate of over fourfold. The high growth ability of this multicellular alga would provide the most effective method for CO2 fixation and biomass production.

Effect of allelochemicals on photosynthetic and antioxidant defense system of Ulva prolifera.

Ulva prolifera is a macroalgae that forms massive blooms, negatively impacting natural communities, aquaculture operations and recreation. The effects of the natural products, eugenol, ß-myrcene, citral and nonanoic acid on the growth rate, antioxidative defense system and photosynthesis of Ulva prolifera were investigated as a possible control strategy for this harmful taxon. Negative effects on growth were observed with all four chemicals, due to the excessive production of reactive oxygen species and oxidative damage to the thalli. However, the response of U. prolifera under the four chemicals stress was different at the cellular level. ß-myrcene, the most effective compound in terms of growth inhibition, induced oxidative stress as shown by the damage of total antioxidant capacity (T-AOC) and the downregulation of the glutathione-ascorbate (GSH-ASA) cycle which inhibited the antioxidative system. This chemical also inhibited photosynthesis and photoprotection mechanisms in U. prolifera, resulting in growth limitation. In contrast, U. prolifera was less affected by the second tested chemical, eugenol, and showed no significant change on photosynthetic efficiency in the presence of the chemical. The inhibition effects of the third and fourth tested chemicals, nonanoic acid and citralon, on growth and on the antioxidant defense system in U. prolifera were inferior. These results provide a potential avenue for controlling green tides in the future.

Impacts of a shallow shipwreck on a coral reef: A case study from St. Brandon's Atoll, Mauritius, Indian Ocean.

Shallow shipwrecks, can have severe ecological and toxicological impacts on coral atolls. In 2012, a tuna longliner ran aground on the reef crest of St Brandon's Atoll, Mauritius, broke up into three pieces which was moved by currents and storms into the lagoon. In the months following the grounding, the coral around the wreck became dead and black. Down-current from the wreck, a dense bloom of filamentous algae (Ulva sp.) attached to coral occurred. To determine the ecological effects of the wreck on the system, the marine biota around the wreck, in the algal bloom, and fish reference zones were counted in 2014. Metal concentrations in reference and affected coral was determined using inductively coupled plasma mass spectrometry (ICP/MS). A pronounced difference was seen in the metal concentration pattern between coral from the wreck- and algal zones, and the coral reference zone. While the wreck zone contained the highest abundance of fish, the fish reference zone had the highest species diversity but with fewer fish. We also counted eleven Critically Endangered hawksbill sea turtles Eretmochelys imbricata and significantly more sea cucumbers in the algal zone than the reference zones. The effects of shipwrecks on coral reefs must be considered a threat over periods of years and should be studied further.

Metagenomic Analysis of the Effect of Enteromorpha prolifera Bloom on Microbial Community and Function in Aquaculture Environment.

Enteromorpha prolifera blooms considerably affected coastal environments in recent years. However, the effects of E. prolifera on microbial ecology and function remained unknown. In this study, metagenomic sequencing was used to investigate the effect of E. prolifera bloom on the microbial communities and functional genes in an aquaculture environment. Results showed that E. prolifera bloom could significantly alter the microbial composition and abundance, and heterotrophic bacteria comprised the major groups in the E. prolifera bloom pond, which was dominated by Actinomycetales and Flavobacteriales. The study indicated that viruses played an important role in shaping the microbial community and diversity during E. prolifera bloom. These viruses affected various dominant microbial taxa (such as Rhodobacteraceae, Synechococcus, and Prochlorococcus), which produced an obvious impact on potential nutrient transformation. Functional annotation analysis indicated that E. prolifera bloom would considerably shift the metabolism function by altering the structure and abundance of the microbial community. E. prolifera bloom pond had the low ability of potential metabolic capabilities of nitrogen, sulfur, and phosphate, whereas promoted gene abundance of genetic information processing. These changes in the microbial community and function could produce serious effect on aquaculture ecosystem.

Ecological effects of Ulva prolifera green tide on bacterial community structure in Qingdao offshore environment.

Ulva prolifera green tide is a serious marine ecological problem in China's coastal areas, with recurrent outbreaks occurring annually during late spring and summer since 2007. Marine bacteria communities are closely linked with important ecological functions in coastal environments. Hence, the succession of bacterial community structures in seawater and sediments during the outbreak and extinction stages of U. prolifera green tide were studied in this paper. The results revealed that: (1) the outbreak of a green tide led to changes in the bacterial community structure and a reduction in the abundance and diversity of the bacterial community in the Qingdao offshore region. This change was more significant in sediments than in seawater. (2) The bacterial assemblage appeared to be dominated by members of Deltaproteobacteria, Bacteroidia and Flavobacteria in sediment, and reductions in the relative abundance of Flavobacteria and Alphaproteobacteria were also observed in seawater after the green tide outbreak. There was also a high similarity in species composition of bacteria in sediment (extinction stage) and seawater (outbreak stage). (3) The outbreak of the U. prolifera affected the abundance of functional bacterial communities in the offshore environment such as SRB (sulfate-reducing bacteria), CFB (Cytophaga-Flavobacter-Bacteroides), heterotrophic bacteria and the potentially algicidal bacteria Alteromonadaceae. Overall, U. prolifera affects aquatic ecosystem secondary production, biogenic geochemical cycles and the growth of other seaweeds in Qingdao offshore areas by driving variations in functional bacterial communities.

Layer-by-layer constructed hyaluronic acid/chitosan multilayers as antifouling and fouling-release coatings.

Polyelectrolyte multilayers (PEMs) consisting of hyaluronic acid (HA) and chitosan (Ch) are extensively studied for biomedical applications and suppress bacterial and protein attachment. Here, we prepared and tested HA/Ch PEMs as marine fouling-release coatings. PEMs were constructed by layer-by-layer assembly using spin coating. The multilayers were crosslinked for enhanced stability in the sea water environment by chemical and thermal treatment. Protein-repelling properties of the crosslinked multilayers were investigated by surface plasmon resonance spectroscopy. The marine antifouling and fouling-release properties were tested against the settlement of zoospores of the green alga Ulva linza and the subsequent development and removal of sporelings. With spin coating and thermal crosslinking, a thick yet homogeneous coating was obtained with antifouling properties against marine algal zoospores indicating the potential of these compounds for application in protective coatings.

Reproductive strategy of the floating alga Ulva prolifera in blooms in the Yellow Sea based on a combination of zoid and chromosome analysis.

Green algal blooms have occurred in the Yellow Sea for 13 consecutive years since 2007. However, little is known about the reproductive strategy of the dominant species Ulva prolifera in the field. In particular, it is not clear whether the floating Ulva species are sporophytes or gametophytes, and if their life history is sexual or asexual. In this study, the life history type was determined based on the size, phototactic response, and flagella number for the zoids in at least two successive generations. In addition, chromosome observations were conducted to distinguish the gametophytes and sporophytes in the floating Ulva species. The results showed that the floating Ulva species were all sporophytes with sexual reproductive patterns, thereby indicating that this Ulva species always maintains vegetative growth from April to June during the early stage of the blooms. In addition, we found that the chromosome numbers were 18 for the diploid sporophytes and nine for the haploid male and female gametophytes. These results provide useful information to help understand the explosive growth of these green algal blooms.

Darkness and low nighttime temperature modulate the growth and photosynthetic performance of Ulva prolifera under lower salinity.

In order to understand how darkness/irradiance and low nighttime temperature might alter physiology of Ulva prolifera under lower salinity conditions, we analyzed the growth rates, water content, superoxide dismutase (SOD) activity, total soluble proteins (SPs) and carbohydrates content at the end of dark and light period under three temperature levels (25-25 °C treatment: 25 °C for day and night; 15-15 °C treatment: 15 °C for day and night; 25-15 °C treatment: 25 °C for day with 15 °C for night) and two salinity conditions (15, 25), meanwhile, the pigment content (chlorophyll a and b), chlorophyll fluorescence and photosynthetic oxygen evolution also were determined during light phase. We found that the U. prolifera showed higher growth rate and SOD activity during dark phase at 25 °C, but this dark-induced increase could not be observed at 15 °C. The reasons for this increase varied, however, maybe not included water content and SPs for no significant difference in water content observed under all the treatments, as well as lower SPs content for dark period aside that at 15 °C and salinity 15. Compared to other two temperature treatments, the thalli grown at 25-15 °C showed higher growth rate and the photosynthetic oxygen evolution rate in light phase under salinity 15 conditions, although the maximum relative electron transport rate (rETRmax) showed higher value under 25 °C treatment. These results indicate that the darkness and the lower nighttime temperature maybe responsible reason for the rapid growth of these green tide algae.

Stable carbon and nitrogen isotopic characterization and tracing nutrient sources of Ulva blooms around Jeju coastal areas.

The present investigation was aimed to characterize the Ulva blooms and to identify the probable sources for Ulva blooms along the Jeju Island coast for pertinent control measures. Algal isotope signatures (δ13C, δ15N and δ18O) and tissue nitrogen and carbon were analyzed to map nutrient sources around the Jeju coastal areas. The algal δ13C values were ranged from -20.52 to -4.39‰, while δ15N and δ18O values ranged from 4.26 to 8.29‰ and 12.80-17.34‰, respectively. Moreover, site-specific significant differences were observed in algal stable isotope (δ13C, δ15N and δ18O) values. The bi-plot (δ15N vs δ18O) diagram indicated four dominant nitrogen sources along the Jeju coast, with 1) soil organic nitrogen mixed with livestock wastes (spring water samples and E), the 2) synthetic fertilizer input (A3 and B2), 3) sewage discharge (D1, D2 and I3) and 4) aquaculture waste (fish farm samples, A4, A5, B1, G and I2). Present findings revealed the different potential nitrogen sources for localized increase of algal growth along the Jeju coast. Finally, the present findings could be used as baseline data for efficient nutrient management to remediate Ulva blooms along Jeju coastal environment.

Uncertainty in the optical remote estimation of the biomass of Ulva prolifera macroalgae using MODIS imagery in the Yellow Sea.

A laboratory experiment was conducted to obtain a floating algae index (FAI) of the floating macroalgae (Ulva prolifera), corresponding to various values of biomass per unit area (BPA). A piecewise empirical model was used to fit the statistical relationships between BPA and FAI, corresponding to FAI ≤ 0.2 (BPA ≤ 1.81kg/m2) and FAI ˃ 0.2 (BPA ˃ 1.81 kg/m2). Spectral mixing derived results show that a linear relationship between FAI and BPA is maintained when the BPA of endmembers is less than 1.81 kg/m2. However, when the BPA of the endmembers exceeds 1.81 kg/m2, there is substantial uncertainty in the optical remote estimation of biomass. Although the MODIS-derived FAI of Ulva prolifera is often less than 0.2, it is very difficult to determine whether the FAI results from low BPA (≤ 1.81kg/m2) of the endmembers, or from a low area ratio including high BPA (˃ 1.81 kg/m2), due to pixel mixing. If it is assumed that the unit biomass distribution of pure endmembers is a standard Gaussian distribution, then the uncertainty in the biomass estimation of Ulva prolifera from MODIS data can be expressed. This results in the uncertainty of ~36% in total biomass estimation, ~43% of which was contributed by a few pixels (10% of total pixels) with high FAI (˃ 0.05). The uncertainty in BPA caused by high FAI (˃ 0.05) pixels is about 7.2 times that for low FAI (≤ 0.05) pixels. In future research, the spatial distribution characteristics of the FAI of pure endmembers need to be considered in order to improve the accuracy of optical remote estimation of floating Ulva prolifera.

The different responses of growth and photosynthesis to NH4+ enrichments between Gracilariopsis lemaneiformis and its epiphytic alga Ulva lactuca grown at elevated atmospheric CO2.

We investigated how elevated CO2 affects the responses of Gracilariopsis lemaneiformis and Ulva lactuca to NH4+ enrichments. All algae were incubated under four nutritional conditions (zero addition, 100, 500, and 2500 µM NH4+), and two CO2 levels (390 ppm and 1000 ppm). The growth, photosynthesis, and soluble protein contents of both species increased under the eutrophication condition (100 µM NH4+). However, the growth and carotenoid contents of the two species declined when NH4+ concentration increased. Under the super eutrophication condition (2500 µM NH4+), all indexes measured in G. lemaneiformis were suppressed, while the growth and photosynthesis in U. lactuca changed indistinctively, both compared with the control. Moreover, under the super eutrophication condition, elevated CO2 reduced the suppression in the growth of G. lemaneiformis, but decreased the growth of U. lactuca. Nonetheless, G. lemaneiformis displayed much lower growth rates than U. lactuca under the super eutrophication and elevated CO2 condition.

Salinity mediates the effects of nitrogen enrichment on the growth, photosynthesis, and biochemical composition of Ulva prolifera.

To study the combined effects of multiple nitrogen (N) sources and salinity on the growth and physiology on macroalgae, we cultured Ulva prolifera under three N levels (N0, 0.1235 mg L-1; N1, 0.6 mg L-1; and N2, 4.4 mg L-1; the ratios were 18:74:8 for NH4-N, NO3-N, and NO2-N, respectively) and three salinity conditions (15, 25, and 35). Then, the growth, pigment content, photosynthetic performance, superoxide dismutase (SOD) activity, and contents of soluble protein and carbohydrates were measured. The results showed the following: (1) Compared to that grown at salinity 25, the growth of U. prolifera decreased under salinity 35, especially under the N0 and N2 levels, but there were no significant effects of salinity 15 under any of the N levels. (2) There were no significant effects of salinity on the chlorophyll a (Chla) content, but compared to the content at salinity 25, the chlorophyll b (Chlb) content was enhanced by salinity 15 and 35; lower ratio values between Chla and carotenoids (Car) occurred under the salinity 25 treatment. Under each salinity condition, the pigments were enhanced by a high N level. (3) A relatively higher salinity level decreased the photosynthetic oxygen evolution rate, while a higher N level increased this value. Compared to the rate at salinity 25, the dark respiration rate (Rd) significantly increased at salinity 15 under the N0 condition. (4) SOD activity was enhanced by a high N level, but no significant effects of salinity were observed. (5) The carbohydrate content was enhanced at salinity 35 under the N0 and N1 levels, and under salinity 15, this value increased with increasing N levels. In conclusion, although the growth of U. prolifera decreased at high N levels under high salinity conditions, a high N level induced an increase in photosynthesis, while no significant decrease in growth occurred. These findings indicate that low salinity and high N levels may be nonnegligible reasons why this species thrives, and low salinity was the better choice when this species was used for wastewater treatment.

Efficient production of poly γ-d-glutamic acid from the bloom-forming green macroalgae, Ulva sp., by Bacillus sp. SJ-10.

Numerous studies on poly γ-d-glutamicacid (γ-PGA) production have investigated terrestrial renewable sources for reducing production costs, but there are no studies using waste marine resources so far. We aimed to develop a cost-effective production method of γ-d-PGA by Bacillus sp. SJ-10 using green macroalgae (Ulva sp.) as a major substrate without hydrolysis pretreatment. The SJ-10 was shown to not only cause immediate tissue degradation of the Ulva membrane but also grew well as a sole substrate. The γ-d-PGA yield was 6.29 ± 0.34 g/L under optimized conditions via the response surface method, and the produced γ-d-PGA had a thermal decomposition temperature of 310°C and molecular weight of 250-1780 kDa. The calculated cost efficiency for the final yield was 32% when compared with complex media. Therefore, the present study provided a strategy for promoting an ecofriendly and cost-effective means to produce γ-d-PGA via a marine renewable resource.

Buoyancy potential of dominant green macroalgal species in the Yellow Sea's green tides, China.

Large-scale green tides caused by Ulva prolifera, occurred for 12 consecutive years in the Yellow Sea of China. To resolve the abrupt shift in species composition between attached and floating macroalgal assemblages, field experiments were conducted from May to July 2017 to quantify the net buoyancy force and compare the floating potential of the common green macroalgae from the red algal seaweed Pyropia yezoensis rafts. At the same time, U. prolifera from different sampling locations were tested to study variable buoyancy of this species and the associated influencing factors. Our results illustrated a stronger positive buoyant force and a proportionally greater buoyancy capacity of U. prolifera, compared to the other co-occurring species. Buoyancy is a dynamic trait and is closely correlated with light intensity, morphology and physiological status. The positive buoyancy of U. prolifera is an important factor that helps explain its predominance in the Yellow Sea's large-scale green tides.

Remote sensing estimation of the biomass of floating Ulva prolifera and analysis of the main factors driving the interannual variability of the biomass in the Yellow Sea.

Since 2007, green tide blooms with Ulva prolifera as the dominant species have occurred every summer in the Yellow Sea. Biomass is a critical parameter used to describe the severity of green tide blooms. In this study, we analyzed the relationships between several indices (normalized difference vegetation index (NDVI), floating algae index (FAI), ratio vegetation index (RVI), enhanced vegetation index (EVI), ocean surface algal bloom index (OSABI), Korea Ocean Satellite Center (KOSC) approach) and the biomass per unit area of Ulva prolifera by using the in situ measurements from a water tank experiment. EVI, NDVI, and FAI showed strong exponential relationships with Ulva prolifera biomass per unit area. In order to apply the relationships to satellite remote sensing data, the impacts of the atmosphere (different aerosol optical depth at 550 nm) and mixed pixels to the relationships were analyzed. The results show that atmosphere has little effect on the relationship between EVI and Ulva prolifera biomass per unit area with R2 = 0.94 and APD (the average percentage deviation) = 19.55% when EVI is calculated from Rrc (Rayleigh-corrected reflectance), and R2 = 0.95 and APD = 17.53% when EVI is calculated from Rtoa (top-of-atmosphere reflectance). Due to the low sensitivity to the atmosphere, the EVI relationship can be directly utilized in the top-of-atmosphere (TOA) reflectance without atmospheric correction. In addition, the EVI was slightly affected by mixed pixels with the APD only increased by ~10%. The EVI relationship was then applied to a long MODIS image time series to obtain the maximal total biomass of floating Ulva prolifera in the Yellow Sea from 2007 to 2016. The results showed that the maximum and minimum total biomass occurred in 2016 (~1.17 million tons) and 2012 (~0.074 million tons), respectively. The main factors that caused the inter-annual biomass variability were analyzed. The total amount of nutrients from Sheyang River which was the largest river on the northern coast of Jiangsu Province, and Porphyra cultivation in the Radial Sand Ridges of Jiangsu Province had both strong correlation with Ulva prolifera total biomass.

Extensive Variations in Diurnal Growth Patterns and Metabolism Among Ulva spp. Strains.

Green macroalgae of the genus Ulva play a key role in coastal ecosystems and are of increasing commercial importance. However, physiological differences between strains and species have yet to be described in detail. Furthermore, the strains of Ulva used in aquaculture usually originate from opportunistic collection in the wild without prior selection of best performing strains. Hence, efforts are required to detect the potential variability in growth and metabolic accumulation between Ulva strains and ultimately select the best performing strains under given environmental conditions. Here, the growth, physiological, and metabolic characteristics of 49 laminar Ulva spp. strains were investigated using a custom-made high-throughput phenotyping platform, enzymatic assays, and gas chromatography-mass spectrometry. We found large natural variation for a wide range of growth and metabolic characteristics, with growth rates varying from 0.09 to 0.37 mg.mg-1d-1 among strains. Ulva spp. possess a unique diurnal growth pattern and primary metabolism compared with land plants, with higher growth rates during the night than during the light period. Starch and sucrose only contributed on average 35% of the carbon required to sustain Ulva's night growth. Nitrates accumulated during the night in Ulva tissues, and nitrate accumulation and consumption was positively correlated with growth. In addition, we identified six amino acids as possible biomarkers for high growth in Ulva The large variability in growth and metabolite accumulation recorded among morphologically similar Ulva strains justifies future efforts in strain selection for increasing biomass, metabolite yields, and nutrient removal in the growing aquaculture industry.