Allelopathic interactions between the opportunistic species Ulva prolifera and the native macroalga Gracilaria lichvoides

Allelopathy, one type of direct plant competition, can be a potent mechanism through which plant communities are structured. The aim of this study was to determine whether allelopathic interactions occur between the opportunistic green tide-forming species Ulva prolifera and the native macroalga Gracilaria lichvoides, both of which were collected from the coastline of East China sea. In laboratory experiments, the presence of G. lichvoides at 1.25 g wet weight L(-1) significantly inhibited growth and photosynthesis of U. prolifera at concentrations of 1.25, 2.50, and 3.75 g wet weight L(-1) (p<0.05) in both semi-continuous co-culture assays and in co-culture assays without nutrient supplementation. In contrast, although U. prolifera had a density effect on G. lichvoides, the differences among treatments were not significant (p>0.05). Culture medium experiments further confirmed that some allelochemicals may be released by both of the tested macroalgae, and these could account for the observed physiological inhibition of growth and photosynthesis. Moreover, the native macroalgae G. lichvoides was a stronger competitor than the opportunistic species U. prolifera. Collectively, the results of the present study represent a significant advance in exploring ecological questions about the effects of green tide blooms on the macroalgal community.     

Effects of macroalgae Ulva pertusa (Chlorophyta) and Gracilaria lemaneiformis (Rhodophyta) on growth of four species of bloom-forming dinoflagellates

The effects of fresh thalli and culture medium filtrates from two species of marine macroalgae, Ulva pertusa Kjellm (Chlorophyta) and Gracilaria lemaneiformis (Bory) Dawson (Rhodophyta), on growth of marine microalgae were investigated in co-culture under controlled laboratory conditions. A selection of microalgal species were used, all being identified as bloom-forming dinoflagellates: Prorocentrum donghaiense Lu sp., Alexandrium tamarense (Lebour) Balech, Amphidinium carterae Hulburt and Scrippsiella trochoide (Stein) Loeblich III. Results showed that the fresh thalli of either U. pertusa or G. lemaneiformis significantly inhibited the microalgal growth, or caused mortality at the end of the experiment. However, the overall effects of the macroalgal culture filtrates on the growth of the dinoflagellates were species-specific (inhibitory, stimulatory or none) for different microalgal species. Results indicated an allelopathic effect of macroalga on the co-cultured dinoflagellate. We then took P. donghaiense as an example to further assess this hypothesis. The present study was carried out under controlled conditions, thereby excluded the fluctuation in light and temperature. Nutrient assays showed that nitrate and phosphate were almost exhausted in G. lemaneiformis co-culture, but remained at enough high levels in U. pertusa co-culture, which were well above the nutrient limitation for the microalgal growth, when all cells of P. donghaiense were killed in the co-culture. Daily f/2 medium enrichment greatly alleviated the growth inhibition on P. donghaiense in G. lemaneiformis co-culture, but could not eliminate it. Other environmental factors, such as carbonate limitation, bacterial presence and the change of pH were also not necessary for the results. We thus concluded that allelopathy was the most possible reason leading to the negative effect of U. pertusa on P. donghaiense, and the combined roles of allelopathy and nutrient competition were essential for the effect of G. lemaneiformis on P. donghaiense.     

Comparative study on mitogenomes of green tide algae

Since 2007, the annual green tide disaster in the Yellow Sea has brought serious economic losses to China. There is no research on the genetic similarities of four constituent species of green tide algae at the genomic level. We previously determined the mitochondrial genomes of Ulva prolifera, Ulva linza and Ulva flexuosa. In the present work, the mitochondrial genome of another green tide (Ulva compressa) was sequenced and analyzed. With the length of 62,311 bp, it contained 29 encoding genes, 26 tRNAs and 10 open reading frames. By comparing these four mitochondrial genomes, we found that U. compressa was quite different from the other three types of Ulva species. However, there were similarities between U. prolifera and U. linza in the number, distribution and homology of open reading frames, evolutionary and codon variation of tRNA, evolutionary relationship and selection pressure of coding genes. Repetitive sequence analysis of simple sequence repeats, tandem repeat and forward repeats further supposed that they have evolved from the same origin. In addition, we directly analyzed gene homologies and translocation of four green tide algae by Mauve alignment. There were gene order rearrangements among them. With fast-evolving genomes, these four green algal mitochondria have both conservatism and variation, thus opening another window for the understanding of origin and evolution of Ulva.     

Tempo-spatial distribution and species diversity of green algae micro-propagules in the Yellow Sea during the large-scale green tide development

From 2008 to 2013, vast green tides mainly composed of Ulva prolifera consecutively invaded the coast of Qingdao (36°06′N, 120°25′E, PR China) in June and July. Previous studies have shown that the early green tides initially formed in the Porphyra yezoensis aquaculture area of the Subei Shoal, southern Yellow Sea. To date, multiple studies have demonstrated that green algae micro-propagules play an important role in the formation of green tides. In this study, we aimed to assess the temporal and spatial distribution of green algae micro-propagules in an extensive area of the Yellow Sea and to determine the species diversity of propagules during the development of the large-scale green tide. We found that the quantity of micro-propagules increased with the free-floating biomass from the initial generation to the development phase of the green tide in mid May. From late May to mid June, the micro-propagule density decreased sharply despite a continuous increase of the floating macroalgae biomass. In addition, our data indicate that the coastal area of the Subei Shoal has always been the distribution center of the micro-propagules, even prior to the large-scale green tide formation. Furthermore, diverse green algae species, including Ulva prolifera, Ulva linza, Ulva flexuosa, Ulva compressa, Ulva pertusa and Blidingia sp., were identified among the micro-propagules in the survey sea area. Finally, we determined that the distribution of U. prolifera micro-propagules is closely related to the floating algal mats and attached macroalgae on Porphyra aquaculture rafts.     

Bioaccumulation of trace metals by the live macroalga Gracilaria lemaneiformis

The macroalga Gracilaria lemaneiformis is an important and commercially valuable renewable resource. It is distributed widely in shallow marine waters but grows mostly on tropical or subtropical coasts. We investigated the accumulation of Cd, Cu, and Pb by live G. lemaneiformis under low concentrations. There was a positive correlation between the organisms’ metal concentrations and exposure concentrations. When exposed to both Cu and Cd, the concentrations of Cu and Cd in G. lemaneiformis were higher than those exposed to solutions of Cu and Cd alone. However, the concentrations of heavy metals in G. lemaneiformis were not markedly different (p?<?0.05) between the treatment groups and the control groups. We analyzed the results with nonlinear curve fitting and employed a two-compartment model to study the accumulation kinetics of heavy metals by G. lemaneiformis. The uptake rate constants and bioconcentration factors (BCFs) of the metals decreased with increased exposure concentration. The theoretical equilibrium concentrations increased significantly with the exposure concentrations. Our results suggested that G. lemaneiformis obviously accumulated heavy metals from seawater. As an important resource for food and pharmaceuticals, G. lemaneiformis should be cultivated in clear seawater.     

Effects of two species of macroalgae—Ulva pertusa and Gracilaria lemaneiformis—on growth of Heterosigma akashiwo (Raphidophyceae)

The effects of fresh thalli, culture filtrate, water-soluble extract and dry powder of two species of macroalgae, Ulva pertusa (Chlorophyta) and Gracilaria lemaneiformis (Rhodophyta), on the growth of a bloom-forming microalga, Heterosigma akashiwo, were studied in co-culture under controlled laboratory conditions. Both fresh thalli and culture filtrate of U. pertusa and G. lemaneiformis, particularly in the form of fresh thalli, significantly inhibited microalgal growth; indeed, the microalga was completely killed during the course of the experiment. A clear concentration-dependent relationship was observed between the initial concentration of fresh thalli (either U. pertusa or G. lemaneiformis) and its inhibitory effect on H. akashiwo. Simultaneous nutrient assays showed that nitrate and phosphate were almost exhausted in G. lemaneiformis fresh thalli co-culture but remained well above nutrient limitation for microalgal growth in U. pertusa co-culture, in which the microalgal cells were completely killed. However, daily f/2 medium repletion would obviously alleviate the growth inhibition in G. lemaneiformis co-culture. Since the present study was carried out under controlled conditions, fluctuations in environmental factors (i.e., light, temperature, carbon limitation, bacterial presence and pH) were limited during the experiment. We thus concluded that allelopathy was the most likely explanation for microalgal growth inhibition in U. pertusa co-culture, while the combined roles of allelopathy and nutrient limitation were responsible for growth inhibition in G. lemaneiformis co-culture. Similarly, macroalgal water-soluble extracts and dry powders affected the co-cultured H. akashiwo greatly, with more obvious effects observed in water-soluble extract co-cultures. A dose-dependent relationship was also observed over the course of the experiment. It can be concluded that macroalgal thalli contain some bioactive compounds. The results of the present study suggest that U. pertusa and G. lemaneiformis, especially in the form of fresh thalli, effectively inhibit the growth of H. akashiwo and could thus be potential candidates for use in the control and mitigation of H. akashiwo blooms.     

Effect of temperature and irradiance on the growth and reproduction of the green macroalga, Chaetomorpha valida (Cladophoraceae, Chlorophyta)

“Green tides” are vast accumulations of unattached green macroalgae associated with eutrophicated marine environments. It had major ecological and economic impacts globally, thus understanding of their detailed conditions was required to inform management decisions. Chaetomorpha valida (Cladophoraceae, Chlorophyta), as a fouling green alga, was found in aquaculture ponds along the coast of Dalian and Rongcheng cities, People’s Republic of China in the past few years. Seasonal abundances of this macroalga in China have presented a significant nuisance and caused great loss to the local aquaculture. Using unialgal cultures, the effect of temperature and irradiance on growth and reproduction were investigated. Healthy growth and reproduction occurred in the range of 17–29°C while the upper lethal limit was at 33°C. A suitable temperature range over 21–29°C and a relatively high irradiance of 108?μmol photons?m^?2?s^?1 were more favorable for growth and reproduction. According to our observations, the diversity of reproductive approaches and wide adaption to temperature can be considered to be key factors that facilitate its excessive growth and colonization in spring and summer. Our results provide a basis for establishing a forecast system and taking remediation measures against seasonal green tides of this macroalga.     

Thermal Acclimation of Respiration and Photosynthesis in the Marine Macroalga Gracilaria lemaneiformis (Gracilariales,Rhodophyta)

The responses of respiration and photosynthesis to temperature fluctuations in marine macroalgae have the potential to significantly affect coastal carbon fluxes and sequestration. In this study, the marine red macroalga Gracilaria lemaneiformis was cultured at three different temperatures (12, 19, and 26°C) and at high‐ and low‐nitrogen (N) availability, to investigate the acclimation potential of respiration and photosynthesis to temperature change. Measurements of respiratory and photosynthetic rates were made at five temperatures (7°C–33°C). An instantaneous change in temperature resulted in a change in the rates of respiration and photosynthesis, and the temperature sensitivities (i.e., the Q₁₀ value) for both the metabolic processes were lower in 26°C‐grown algae than 12°C‐ or 19°C‐grown algae. Both respiration and photosynthesis acclimated to long‐term changes in temperature, irrespective of the N availability under which the algae were grown; respiration displayed strong acclimation, whereas photosynthesis only exhibited a partial acclimation response to changing growth temperatures. The ratio of respiration to gross photosynthesis was higher in 12°C‐grown algae, but displayed little difference between the algae grown at 19°C and 26°C. We propose that it is unlikely that respiration in G. lemaneiformis would increase significantly with global warming, although photosynthesis would increase at moderately elevated temperatures.     

Effects of temperature and irradiance on photosynthesis and growth of a green-tide-forming species (<Emphasis Type="Italic">Ulva linza</Emphasis>) in the Yellow Sea

Samples of the massive drifting green alga, Ulva linza, were collected from the coastal waters of the Yellow Sea, southwest of Korea, in early July 2009, and cultured under laboratory conditions. The effects of various temperature (10–30°C) and irradiance levels (0–1,000 μmol photons m⁻² s⁻¹) on photosynthesis, growth, and tissue nutrient content of U. linza were investigated in laboratory for both individuals of the late-stage vegetation (LSV) and the early-stage vegetation (ESV). After 1 h acclimation to various irradiance and temperature conditions, maximum gross photosynthetic rate of ESV was significantly higher than those of LSV. In the long-term (7-d) acclimation experiments to various irradiance and temperature levels, gross photosynthetic rates of ESV individuals were also significantly higher than those of LSV individuals. High photosynthetic rate of ESV individuals induced increase in mass of about 60% over the growth saturation irradiance (136 μmol photons m⁻² s⁻¹) and about 20% under low temperature conditions (10–15°C) during 7-d. The gross photosynthesis of LSV individuals was low when examined under temperature and irradiance conditions that were optimum for ESV growth. Consequently, free-floating U. linza exhibits cellular senescence beginning in early July in the Yellow Sea, and green tides formed by this species cannot be maintained beyond this time in the open sea. However, we expect that U. linza can proliferate quickly after settlement on new coastal habitats of the Yellow Sea because of the high tissue nitrogen utilization for photosynthesis in ESV, which is formed by germination of reproductive cells.     

Quantification of Selected Endogenous Hydroxy-oxylipins from Tropical Marine Macroalgae

The present study investigated the contents of hydroxy-oxylipins hydroxyoctadecadienoic acids (HODEs), hydroxyoctadecatrienoic acids (HOTrEs), and hydroxyeicosatetraenoic acids (HETEs) in 40 macroalgae belonging to the Chlorophyceae, Rhodophyceae and, Phaeophyceae. The hydroxy-oxylipin content was low and ranged from 0.14?±?0.012 ng/g (Codium dwarkense) to 8,161.9?±?253 ng/g (Chaetomorpha linum) among the Chlorophyceae, 345.4?±?56.8 ng/g (Scytosiphon lomentaria) to 2,574.5?±?155.5 ng/g (Stoechospermum marginatum) among the Phaeophyceae, and 19.4?±?2.2 ng/g (Laurencia cruciata) to 1,753.1?±?268.2 ng/g in Gracilaria corticata v. folifera) among the Rhodophyceae on fresh weight basis (p?≤?0.01). The concentrations of C18-oxylipins were greater than C20-oxylipins in all the investigated macroalgae, except forUlva linza, Codium sursum, Dictyopteris deliculata, S. marginatum, Sargassum tenerrimum, Gracilaria spp. (except G. textorii), Rhodymenia sonderi, and Odonthalia veravalensis.The macroalgal species rich in HODEs, HOTrEs, and HETEs were segregated using principal component analysis. The red macroalgae showed the highest contents of HETEs, followed by brown and green macroalgae in consistent with their PUFA profiles. The relative contents of isomeric forms of oxylipins displayed the species-specific positional selectivity of lipoxygenase (LOX) enzyme in macroalgae. All the species exhibited 13-LOX specificity for linoleic acid analogous of higher plants, while 21 out of 40 species showed 9-LOX selectivity for the oxygenation of α-linolenic acid. No trend was observed for the oxygenation of arachidonic acid in macroalgae, except for in the Halymeniales, Ceramiales (except L. cruciata), and Corallinales. This study infers that LOX products, octadecanoids and eicosanoids, described in macroalgal taxa were similar to those of higher plants and mammals, respectively, and thus can be utilized as an alternative source of chemically synthesized oxylipin analogues in therapeutics, cosmetics, and nutritional oil supplements.     

Physiological effects of nitrogen deficiency and recovery on the macroalga Gracilariopsis lemaneiformis (Rhodophyta)

Algal metabolites are the most promising feedstocks for bio‐energy production. Gracilariopsis lemaneiformis seems to be a good candidate red alga for polysaccharide production, especially relating to the agar production industry. Nitrogen deficiency is an efficient environmental pressure used to increase the accumulation of metabolites in algae. However, there are no studies on the physiological effects of G. lemaneiformis in response to nitrogen deficiency and its subsequent recovery. Here we integrated physiological data with molecular studies to explore the response strategy of G. lemaneiformis under nitrogen deficiency and recovery. Physiological measurements indicated that amino acids and protein biosynthesis were decreased, while endogenous NH₄⁺ and soluble polysaccharides levels were increased under nitrogen stress. The expression of key genes involved in these pathways further suggested that G. lemaneiformis responded to nitrogen stress through up‐regulation or down‐regulation of genes related to nitrogen metabolism, and increased levels of endogenous NH₄⁺ to complement the deficiency of exogenous nitrogen. Consistent with the highest accumulation of soluble polysaccharides, the gene encoding UDP‐glucose pyrophosphorylase, a molecular marker used to evaluate agar content, was dramatically up‐regulated more than 4‐fold compared to the relative expression of actin after 4 d of nitrogen recovery. The present data provide important information on the mechanisms of nutrient balance in macroalgae.     

Seasonal variation of dominant free-floating and attached Ulva species in Rudong coastal area,China

In this paper, species compositions and seasonal variations of attached Ulva species on Porphyra aquaculture rafts and free floating Ulva species at Rudong coastal area, Jiangsu Province of China were investigated during 2010–2011. Based on the sequences analysis of nuclear-encoded ITS (including 5.8S rDNA regions) and 5S rDNA spacer regions, dominant species of both attached and free-floating Ulva samples were identified as Ulva compressa, Ulva linza, Ulva prolifera and Ulva flexuosa. Phylogenetic tree based on sequences of ITS and 5S rDNA spacer regions for attached and free-floating Ulva species was constructed, respectively. Species compositions of the Ulva population attached on aquaculture rafts varied with seasons, and U. prolifera was only found on aquaculture rafts in March 2011 during the 2010–2011 Porphyra yezoensis cultivation season, which had the same sequences of ITS and 5S rDNA spacer regions as that of the dominant species bloomed in the Yellow Sea of China in 2008. Dominant species of the free-floating Ulva population at the early stage of the green tide were U. compressa, U. flexuosa, and U. linza. Free-floating U. prolifera appeared in the middle of May, 2011. ITS sequence similarity rates of U. compressa and U. flexuosa between the attached and free-floating species were 100%. And ITS and 5S rDNA spacer sequences of the attached and the free-floating U. prolifera population also showed no differences. Further study showed that there were two types of free-floating U. prolifera population (Type 5S-A and Type 5S-B) based on 5S rDNA spacer sequences. The present study would provide some useful information for clarifying the outbreak mechanism of green tides occurred in the Yellow Sea, China.     

Effects of elevated nutrients and CO2 emission scenarios on three coral reef macroalgae

Coral reef macroalgae are expected to thrive in the future under conditions that are deleterious to the health of reef-building corals. Here we examined how macroalgae would be affected by exposure to future CO₂ emission scenarios (pCO₂ and temperature), enriched nutrients and combinations of both. The species tested, Laurencia intricata (Rhodophyta), Turbinaria ornata and Chnoospora implexa (both Phaeophyceae), have active carbon-concentrating mechanisms but responded differently to the treatments. L. intricata showed high mortality under nutrient enriched RCP4.5 (“reduced” CO₂ emission) and RCP8.5 (“business-as-usual” CO₂ emission) and grew best under pre-industrial (PI) conditions, where it could take up carbon using external carbonic anhydrase combined, potentially, with proton extrusion. T. ornata’s growth rate showed a trend for reduction under RCP8.5 but was unaffected by nutrient enrichment. In C. implexa, highest growth was observed under PI conditions, but highest net photosynthesis occurred under RCP8.5, suggesting that under RCP8.5, carbon is stored and respired at greater rates while it is directed to growth under PI conditions. None of the species showed growth enhancement under future scenarios, nutrient enrichment or combinations of both. This leads to the conclusion that under such conditions these species are unlikely to pose an increasing threat to coral reefs.     

Growth characteristics and reproductive capability of green tide algae in Rudong coast, China

Since 2007, green tides have occurred along the coast of the Yellow Sea, China. The green tide extended to 50,000 km² (floating area) within 2–3 months and the calculated covering area was about 400 km² in 2010. These facts implied that the growth and reproduction of the dominant species Ulva prolifera were stimulated. We observed that 1 cm² blades (single layer) released 2.84–6.62?×?10⁶ spores or 1.14–2.65?×?10⁷ gametes and that 91.6–96.4 % of them germinated into younger seedlings. This means that, in theory, 1 g (fresh weight) of blades was able to produce about 2.8?×?10⁸–2.7?×?10⁹ new younger seedlings. From 2009 to 2011, the growth rate of green tide algae was measured in situ in enclosure experiments in Rudong coast, Jiangsu Province and the growth curve of the algae was divided into four phases: lag phase, accelerated phase, stationary phase, and decline phase. Usually, the average daily specific relative growth rate was about 23.2–23.6 % d^?1 for a whole growth period, and it reached up to 56.2 % d^?1 in the accelerated phase. Correspondingly, the morphology of green tide algae in enclosures also showed periodic variation as follows: blades presented new filamentous branches from old thallus in the lag phase, longer filamentous branches in the accelerated phase, tubular and cystic blades in the stationary phase, and folded blades in the decline stage. Those studies may be useful for understanding the green tide blooming mechanism.     

MIPS: A Calmodulin-Binding Protein of Gracilaria Lemaneiformis Under Heat Shock

To study the Ca²⁺/Calmodulin (CaM) signal transduction pathway of Gracilaria lemaneiformis under heat stress, myo-inositol-1-phosphate synthase (MIPS), a calmodulin-binding protein, was isolated using the yeast two-hybrid system. cDNA and DNA sequences of mips were cloned from G. lemaneiformis by using 5′RACE and genome walking procedures. The MIPS DNA sequence was 2,067 nucleotides long, containing an open reading frame (ORF) of 1,623 nucleotides with no intron. The mips ORF was predicted to encode 540 amino acids, which included the conserved MIPS domain and was 61–67 % similar to that of other species. After analyzing the amino acid sequence of MIPS, the CaM-Binding Domain (CaMBD) was inferred to be at a site spanning from amino acid 212 to amino acid 236. The yeast two-hybrid results proved that MIPS can interact with CaM and that MIPS is a type of calmodulin-binding protein. Next, the expression of CaM and MIPS in wild-type G. lemaneiformis and a heat-tolerant G. lemaneiformis cultivar, "981," were analyzed using real-time PCR under a heat shock of 32 °C. The expression level displayed a cyclical upward trend. Compared with wild type, the CaM expression levels of cultivar 981 were higher, which might directly relate to its resistance to high temperatures. This paper indicates that MIPS and CaM may play important roles in the high-temperature resistance of G. lemaneiformis.     

Effects of deep seawater medium on growth and amino acid profile of a sterile Ulva pertusa Kjellman (Ulvaceae, Chlorophyta)

The potential use of deep seawater (DSW) for the cultivation of macroalgae was evaluated by analysis of the growth rate of thalli (by surface area), photosynthetic pigment content, and total free amino acid content of sterile Ulva pertusa cultured in the control medium (one-fifth strength Provasoli’s enriched seawater (PES) medium), surface seawater (SSW) medium and also seawater medium collected from 400 m depth (400-DSW) and 1,200 m depth (1,200-DSW). The growth rate of the sterile U. pertusa grown in 400-DSW and 1,200-DSW was similar to that of thalli cultured in the one-fifth strength PES medium that had previously been shown to be the optimum concentration for the cultivation of Ulva. Photosynthetic pigment content in the control medium, 400-DSW- and 1,200-DSW-cultured thalli was 1.7–2.0 times higher than that of the SSW-cultured thalli. The total free amino acids content of the control medium and SSW-cultivated thalli was 264?±?64 and 120?±?39 mg per 100 g dry weight, whereas the 400-DSW- and 1,200-DSW-cultivated thalli had significantly more amino acids (342?±?69–327?±?64 mg per 100 g dry weight). The results of this study indicate that DSW has a high potential for cultivation of edible macroalgae without any supply of extra nutrient until it grows to harvest size.     

Artificial substrates enhance non-native macroalga and N2 production

The introduction of non-native species occurs within a context of other anthropogenic impacts: thus a holistic approach is needed to understand interactive effects. Installation of shoreline protection structures is increasing in response to rising sea levels and increasing frequency of intense storms. Shoreline hardening structures can facilitate establishment of non-native species with multiple potential consequences. We measured abundances of both native and non-native Codium, a green macroalga, on natural hard substrates (oyster reefs) and artificial substrates (bulkheads and revetments) in two estuaries and assessed the effects of each Codium species on nitrogen fixation and net N₂ fluxes. Native C. decorticatum was the most abundant (86 %) Codium on oyster reefs, while exotic C. fragile dominated (99 %) artificial substrates. N₂ production via denitrification was greater than nitrogen fixation for both species and the net N₂ production associated with non-native C. fragile was greater than with native C. decorticatum. Comparing our results with surveys conducted in the 1940s before C. fragile had invaded, indicates that non-native Codium has outcompeted native Codium on artificial substrates, but that natural substrate provided by oyster reefs remains a refuge for native Codium. Although shoreline hardening with artificial structures can reduce ecosystem services provided by coastal marsh and other habitats, an unexpected beneficial consequence was the N₂ production associated with the non-native Codium, which has the potential to mitigate anthropogenic nutrient loading. Our results illustrate that the interaction between multiple anthropogenic impacts can be positive, and indicate that non-ephemeral macroalgae could be an overlooked component in nitrogen removal from marine ecosystems by enhancing denitrification.