Stabilisation of ‘Green tides’ Ulva by a method of composting with a view to pollution limitation

85 000 m³ of Ulva sp. is harvested each year in Brittany (France) during a 3–4-month season. Spreading is often impossible and the alga does not keep for more than a few days. Storage results in objectionable odours and release of a liquified fraction. Such pollution can be suppressed by composting the alga with the lowest possible quantity of ligno-cellulosic substrates. This results in the stabilisation of the seaweed, and its use can thereby be delayed. This system of processing the seaweed decreases the cost of treatment by composting and the product can be used either as a substrate for later composting or as an organic enhancing and fertilising agent. (*author for correspondence)     

Changes in biomass and botanical composition of beach-cast seaweeds in a disturbed coastal area from Argentine Patagonia

Trends in wrack composition and biomass, and its relationship with the anthropogenic impact were studied along a coastal area in Nuevo Gulf (south Patagonia) in front of Puerto Madryn city. Beach-cast macroalgae composition was sampled from 1992 to 1999 in the Puerto Madryn beaches and in several other nearby beaches in 1993, 1996 and 1998. Historical information was based on local knowledge and observations reported by marine biologists who worked in the area. The botanical composition of the beach-cast macroalgae in Puerto Madryn indicates a succession in the dominance from Codium spp. to Ulva spp. during the 1990s and from Ulva to Undaria pinnatifida since 1998, accompanied by a significant decrease in biomass of Gracilaria gracilis and Macrocystis pyrifera. The increase of the opportunist species such as Ulva may be supported by the continuous delivery of waste waters into the Nuevo Gulf while the dominance of U. pinnatifida may be associated with port activities. During the sampling period the highest wrack biomass values were recorded in spring and summer. The beach-cast seaweed biomass harvested by the municipality of Puerto Madryn during beach cleaning operation, ranged between 2500 and 12000 t year^–1 ( 200 and 960 t dry weight). Wrack harvesting produces an environmental impact by removing sand from the beach and affecting coastal communities. Composting of wrack is proposed as one of the environmental alternatives to land disposal.     

Utilisation of Ulva rigida biomass in the Venice Lagoon (Italy): biotransformation in compost

In the Venice Lagoon (Italy), about 10⁶ t (wet weight) of Ulva rigida biomass are produced annually as one of the major results of eutrophication. Harvests have been initiated to reduce negative impacts of this biomass, however, due to the high costs of such effort, only 40 000 t yr^-1 are currently being collected. At the moment, biotransformation into compost seems to be the only feasible technology for utilising large quantities of Ulva biomass. We describe and discuss here a successful composting strategy together with the chemico-physical and microbiological characteristics of the resultant composts. Our composting experiments were conducted at a scale of 20 t. The composting technology utilises large proportions (70–90%) of Ulva biomass and results in a valuable, high-quality end product (compost and compost-based products). This process and the resulting products represents a relatively simple way of utilising the Ulva biomass produced annually in the Venice Lagoon.     

Commercial potential of seaweeds from St Lawrence Island,Alaska

This study assessed the possibility of using drift and subtidal seaweeds from St Lawrence Island, Alaska (lat. 63°N) for sale by the native population after simple processing. Over 125 km of coastline were surveyed for distribution of both drift and subtidal seaweeds. Drift seaweed wet weight densities ranged from 0.2 to over 9 kg m⁻², with an average of over 4 t km⁻¹ in the areas sampled. Attached, benthic seaweed densities ranged from 0.15 to 0.32 kg m⁻². Thirty and 35% of the biomass was composed of Agarum cribrosum and species of Laminaria, respectively, both as drift and as benthic seaweed. Data from tagged Laminaria indicated that growth rates were relatively slow for most of the year. The drift seaweed resource on the coasts south and west of the city of Gambell appeared to have good potential for a small-scale commercial harvest. (*author for correspondence)     

Growth of the seaweed Ulva rigida C. Agardh in relation to biomass densities, internal nutrient pools and external nutrient supply in the Sacca di Goro lagoon (Northern Italy)

Growth of the seaweed Ulva rigida C. Agardh was investigated in relation to biomass densities, internal nutrient pools and external nutrient supply. Research was carried out from 23 March to 5 July 1994 in the Sacca di Goro (Po Delta, Northern Italy), whose south-eastern part was covered by extensive mats of Ulva rigida. Two types of field experiments were conducted by incubating Ulva thalli inside large cages. In the first experiment, beginning on 23 March, 100 g of wet thalli were placed into the cages, allowed to grow for two weeks, then collected and replaced. This procedure was repeated 8 times over the study period. In the second experiment, Ulva thalli were left inside the cages and collected at selected time intervals (14, 27, 41, 64 and 76 days) in order to simulate the effects of increased density on growth and nutrient storage.We recorded specific growth rates (NGR) ranging from 0.025 to 0.081 d^–1 for a period up to two months in the repeated short-term experiments performed at relatively low initial algal densities (300–500 g AFDW m^–3). These NGR resulted significantly related to dissolved inorganic nitrogen (DIN) in the water column. Tissue concentrations of total Kjeldahl nitrogen (TN) were almost constant, while extractable nitrate decreased in a similar manner to DIN in the water column. Total phosphorus showed considerable variation, probably linked to pulsed freshwater inflow.In the long-term incubation experiment, NGR of Ulva was inversely related to density. Internal concentrations of both total P and TN reached maximum values after one month; thereafter P concentration remained almost constant, while TN decreased below 2% w/w (by dry weight). The TN decrease was also accompanied by an abrupt decrease in nitrate tissue concentration. The biomass incubated over the two month period suffered a progressive N limitation as shown by a decreasing NY ratio (49.4 to 14.6). The reciprocal control of Ulva against biogeochemical environment and viceversa is a key factor in explaining both resource competition and successional stages in primary producer communities dominated by Ulva. However, when the biomass exceeds a critical threshold level, approximately 1 kg AFDW m^–3, the macroalgal community switches from active production to rapid decomposition, probably as a result of selfshading, biomass density and development of anaerobic conditions within the macroalgal beds.     

The role of Ulva spp. as a temporary nutrient sink in a coastal lagoon with oyster cultivation and upwelling influence

Bahía San Quintín is a coastal lagoon with large Ulva biomass and upwelling influence. Previous observations suggest that Ulva has increased recently as a result of oyster cultivation. To evaluate the possible role of Ulva as a temporary nutrient sink, biomass and tissue C, N, and P were determined seasonally. Maximum biomass was present during spring and early summer (1,413–1,160 t (dry)) covering about 400 ha. However, the biomass decreased to 35 t (dry) by winter. The mean annual percentage of Ulva C, N, and P was 28%, 2%, and 0.14%, respectively. This study shows that Ulva can store up to 28 t of N and 2 t of P in Bahía San Quintín during spring–summer. Ulva may be displacing the seagrass Zostera marina subtidal beds. A partial removal of the seaweed would reduce the risk of eutrophication and the displacement of eelgrass beds.     

Growing Ulva (Chlorophyta) in integrated systems as a commercial crop for abalone feed in South Africa: a SWOT analysis

Over 1,000t fresh weight of Ulva was cultivated on South African abalone farms in 2007, primarily for feed, but in one case to allow partial re-circulation by nutrient removal. The potential of this new commercial crop is analysed. Material is collected from natural free-floating populations and at least four species are commonly grown, with different ecophysiological characteristics. A brown epiphyte, Myrionema strangulans, causes a disease of the Ulva, which is currently managed by farmers by re-stocking. The main potential threat is that some farmers are wary of integrated systems, fearing they may promote disease in abalone, although Ulva has been grown in abalone effluent and fed back to the abalone on one farm for 6 years without adverse effects. Opportunities exist for the expansion of Ulva cultivation via further spread of the abalone industry, the inclusion of seaweed raceways in proposed fish-farming activities, or the potential for the inclusion of high quality cultivated Ulva as a constituent in aquafeed. A conservative estimate of production over a full year from these raceway systems was 26.1 g dw m⁻² day⁻¹ (2006) and 19.7 g dw m⁻² day⁻¹ (2007), similar to maximum figures for total annual microalgal biomass production in outdoor systems.     

Integrated seaweed cultivation on an abalone farm in South Africa

Land-based abalone aquaculture in South Africa, based on the local species Haliotis midae, started in the early 1990s and has grown rapidly in the last decade, with 13 commercial farms now producing over 850 t per annum. Over 6,000 t per annum of kelp Ecklonia maxima are now harvested for this purpose, and some kelp beds are reaching maximum sustainable limits. Research into seaweed aquaculture as feed (Ulva and some Gracilaria) for abalone started in the late 1990s on the southeast coast (where there are no kelp beds) using abalone waste water. A growing body of evidence suggests that a mixed diet of kelp plus other seaweeds can give growth rates at least as good as compound feed, and can improve abalone quality and reduce parasite loads. A pilot scale Ulva lactuca and abalone integrated recirculation unit using 25% recirculation was designed and built on the south west coast of South Africa using one 12,000-L abalone tank containing 13,200 15 ± 2.5 g abalone, connected to two 3,000-L seaweed tanks containing an initial starting biomass of 10 kg of seaweed, replicated 3 times. In an 18-month period, there were no significant differences in abalone health or growth rates, sediment build up and composition, mobile macro fauna densities and species between the recirculation or the flow-through units. Transfer of oxygen generated by the seaweeds to the abalone tanks was poor, resulting in the recirculated abalone tanks having lower (33%) dissolved oxygen concentrations than a comparable flow-through abalone unit. Seaweed nutrient content and specific growth rates in the units were comparable to seaweeds cultivated in fertilized effluent (SGR = 3.2 ± 3.4%.day⁻¹; Yield = 0.2 ± 0.19 kg.m².day⁻¹). Indications were that at this low recirculation ratio the seaweeds in the units were nutrient limited and that there were no negative effects to the abalone being cultivated in such a recirculation unit at this recirculation ratio.     

Nutrient and iron limitation to <Emphasis Type="Italic">Ulva</Emphasis> blooms in a eutrophic coastal lagoon (Sacca di Goro,Italy)

Growth patterns and bloom formation of the green seaweed Ulva rigida were analysed in the eutrophic Sacca di Goro lagoon (Po River Delta, Italy). Variations of standing biomasses and elemental composition of Ulva were analysed through an annual cycle with respect to nitrogen, phosphorus and iron. Growth rates, nutrient and iron uptake and nitrate storage by macroalgal thalli were also assessed with field experiments during the formation of a spring bloom. The control of Ulva growth and the bloom formation depended on multiple factors, especially on nitrogen availability and iron deficiency. In the nitrate rich waters of the Sacca di Goro lagoon, nitrate accumulation in Ulva thalli was inversely related with Fe uptake, indicating an influence of Fe limitation on N acquisition. Since length and magnitude of nitrate luxury uptake are inversely related to the size of the intracellular nitrate pools, in nitrate rich waters the fast growing Ulva may face risk of N-limitation not only when exposed to low N concentrations or at high biomass levels, but also when exposed to pulsed dissolved nitrate concentrations at low iron availability. The potential Fe limitation could be affected by processes controlled by geochemical reactions and by macroalgal growth and decomposition. Both Fe oxidation during the active macroalgal growth and the formation of insoluble FeS and FeS₂ during bloom collapse can result in a drastic decrease of soluble iron. Thus, a potential limitation of Fe to macroalgae can occur, determining positive feedbacks and potentially controlling the extent of bloom development and persistence.     

Bioactive compounds in seaweed: functional food applications and legislation

Seaweed is more than the wrap that keeps rice together in sushi. Seaweed biomass is already used for a wide range of other products in food, including stabilising agents. Biorefineries with seaweed as feedstock are attracting worldwide interest and include low-volume, high value-added products and vice versa. Scientific research on bioactive compounds in seaweed usually takes place on just a few species and compounds. This paper reviews worldwide research on bioactive compounds, mainly of nine genera or species of seaweed, which are also available in European temperate Atlantic waters, i.e. Laminaria sp., Fucus sp., Ascophyllum nodosum, Chondrus crispus, Porphyra sp., Ulva sp., Sargassum sp., Gracilaria sp. and Palmaria palmata. In addition, Undaria pinnatifida is included in this review as this is globally one of the most commonly produced, investigated and available species. Fewer examples of other species abundant worldwide have also been included. This review will supply fundamental information for biorefineries in Atlantic Europe using seaweed as feedstock. Preliminary selection of one or several candidate seaweed species will be possible based on the summary tables and previous research described in this review. This applies either to the choice of high value-added bioactive products to be exploited in an available species or to the choice of seaweed species when a bioactive compound is desired. Data are presented in tables with species, effect and test organism (if present) with examples of uses to enhance comparisons. In addition, scientific experiments performed on seaweed used as animal feed are presented, and EU, US and Japanese legislation on functional foods is reviewed.     

Effect of grazing by isopods and amphipods on growth of Ulva spp. (Chlorophyta)

Eutrophication of shallow coastal waters often leads to blooms of macroalgae. Grazing by crustaceans, such as amphipods and isopods, can reduce macroalgal biomass accumulation. At the same time, growth of the macroalgae can be stimulated by epiphyte removal. The role of grazing by isopods and amphipods on Ulva spp. biomass development was investigated in the Veerse Meer, a brackish lagoon situated in the southwest Netherlands. Exclusion of grazing in the field did not stimulate Ulva spp. growth. In fact, growth rates were higher in exclosures that allowed grazers to enter. Edibility tests identified the amphipod Gammarus locusta, and the isopods Idotea chelipes and Sphaeroma hookeri as potential grazers on Ulva spp. However, when epiphytic diatoms were present on the Ulva spp. thalli, Gammarus and Sphaeroma grazed on ephiphytes and not on Ulva tissue. Only Idotea continued to graze on Ulva spp. A laboratory growth experiment revealed a positive effect of Gammarus presence on Ulva spp. growth, probably caused by preferential removal of epiphytic diatoms from the Ulva spp. thalli. The growth stimulation by epiphyte removing grazers such as Gammarus may explain the higher growth rates in the presence of grazers observed in the field. When determining the potential role of invertebrate grazers in controlling macroalgal biomass accumulation, it is important to include an assessment of the epiphyte abundance on the macroalgae, as preferential removal of epiphytes may stimulate growth and thus have the opposite effect.     

Anaerobic digestion of Ulva sp. 1. Relationship between Ulva composition and methanisation

Ulva often represents the main component of mass algal growths, and its composition and degradability make it a relatively good methanisation substrate. In ‘green tides’ Ulva sp. from Brittany, the low content oflignin-type components (polyphloroglucinols: 1.3% dry weight), and the large hemicellulosic fraction (9% dry weight) favour the substrate's accessibility to enzymes. Anaerobic degradation with a batch orcompletely stirred system is technically possible. However, the methane yield reached only 0.20 m³ kg⁻¹ volatile solids and the epuration rate 50% volatile solids in experiments in batch or completely stirred reactors. More generally, methanisation comes up against various practical obstacles: seasonal growth of Ulva, low density of alga in suspension for loading the digester, high S concentration leading to the production of a biogas with a high H2S content, and, finally, the existence of a refractory or slowly degradable part, which requires a compromise between productivity and biological yield. This revised version was published online in August 2006 with corrections to the Cover Date.     

Dissolved oxygen and nutrient budgets in a phytotreatment pond colonised by <Emphasis Type="Italic">Ulva</Emphasis> spp.

Net daily budgets of dissolved oxygen (O₂), dissolved inorganic carbon (DIC), dissolved inorganic nitrogen (DIN = NH₄⁺+NO₂⁻+NO₃⁻) and soluble reactive phosphorus (SRP) were determined in a pond colonised by Ulva spp. This pond received wastewater from a land-based fish farm and was used as a phytotreatment plant. Three consecutive 24-h cycles of measurements were performed with 8–14 samplings per day. Water samples were collected at the inlet and outlet of the pond and budgets were estimated from differences between inlet and outlet loadings. The first cycle was started when Ulva biomass was 8 kg m⁻², as wet weight. The second cycle was performed after the harvest of ~20% of the macroalgal biomass and the third after the harvest of another ~20% of the remaining biomass. Ulva removal was very fast (<1 h) and samplings for cycles 2 and 3 were started two hours after harvesting, so that the whole experiment lasted ~80 h. When Ulva biomass was at its maximum, the aquatic system was heterotrophic with an O₂ demand of 519 mol d⁻¹ and a net regeneration of DIC (2686 mol d⁻¹), NH₄⁺ (49 mol d⁻¹) and SRP (2.5 mol d⁻¹). The DIC to O₂ ratio was an indicator of persistent anaerobic metabolism. Following the first harvest intervention, this system displayed a prompt response and shifted toward a lower O₂ demand (from −519 to −13 mol d⁻¹), with a lesser regeneration degree of NH₄⁺ (11.4 mol d⁻¹) and DIC (1066 mol d⁻¹). After the second Ulva removal the net budget of SRP became negative (−1.0 mol d⁻¹). By integrating these results over the three days cycle we estimated that in order to operate an efficient nutrient control and maintain macroalgal mats in a healthy status the optimal Ulva biomass should be well below ~4 kg m⁻² as wet weight. Above this threshold, self-limitation would render most of the algal mat unable to exploit light and nutrients. An efficient removal of nitrogen and phosphorus could be attained through the management of macroalgal biomass only with an optimisation of recipient surface to nutrient loading ratio.     

Assessment of the nutritional value of Capsosiphon fulvescens (Chlorophyta): developing a new species of marine macroalgae for cultivation in Korea

The nutritive value of Capsosiphon fulvescens (C. Agardh) Setchell et Gardner, a new developing species for cultivation of marine macroalgae in Korea, was assessed by comparison with common edible green seaweed Ulva prolifera (Oeder) J. Agardh collected from Korea and Japan, based on analysis of its chemical composition. The contents of protein and of total, essential, and free amino acids of C. fulvescens were significantly higher than those of U. prolifera, whereas those of lipid, carbohydrate, and total dietary fibres were significantly lower. There were no significant differences in the moisture content between Capsosiphon and Ulva species. The main minerals of Capsosiphon and Ulva are Na, Mg, K, Ca, and Al, making up approximately 97–99 % of the total minerals. The contents of Na and V of Capsosiphon were significantly greater than those of Ulva, while those of Mn and Sr were significantly lower in Capsosiphon. The contents of retinal (27.8 μg g⁻¹ dry wt) and ascorbic acid (0.28 mg g⁻¹ dry wt) of Capsosiphon were significantly higher than those of Ulva, but the contents of cobalamin were lower. These results suggest that C. fulvescens has greater potential to be used as human food and as an ingredient in formulated food.     

A sixth‐level habitat cascade increases biodiversity in an intertidal estuary

Many studies have documented habitat cascades where two co‐occurring habitat‐forming species control biodiversity. However, more than two habitat‐formers could theoretically co‐occur. We here documented a sixth‐level habitat cascade from the Avon‐Heathcote Estuary, New Zealand, by correlating counts of attached inhabitants to the size and accumulated biomass of their biogenic hosts. These data revealed predictable sequences of habitat‐formation (=attachment space). First, the bivalve Austrovenus provided habitat for green seaweeds (Ulva) that provided habitat for trochid snails in a typical estuarine habitat cascade. However, the trochids also provided habitat for the nonnative bryozoan Conopeum that provided habitat for the red seaweed Gigartina that provided habitat for more trochids, thereby resetting the sequence of the habitat cascade, theoretically in perpetuity. Austrovenus is here the basal habitat‐former that controls this “long” cascade. The strength of facilitation increased with seaweed frond size, accumulated seaweed biomass, accumulated shell biomass but less with shell size. We also found that Ulva attached to all habitat‐formers, trochids attached to Ulva and Gigartina, and Conopeum and Gigartina predominately attached to trochids. These “affinities” for different habitat‐forming species probably reflect species‐specific traits of juveniles and adults. Finally, manipulative experiments confirmed that the amount of seaweed and trochids was important and consistent regulators of the habitat cascade in different estuarine environments. We also interpreted this cascade as a habitat‐formation network that describes the likelihood of an inhabitant being found attached to a specific habitat‐former. We conclude that the strength of the cascade increased with the amount of higher‐order habitat‐formers, with differences in form and function between higher and lower‐order habitat‐formers, and with the affinity of inhabitants for higher‐order habitat‐formers. We suggest that long habitat cascades are common where species traits allow for physical attachment to other species, such as in marine benthic systems and old forest.     

BIOMASS AND DYNAMICS OF GROWTH OF ULVA SPECIES IN PALMONES RIVER ESTUARY1

During the last decade, the Palmones River estuary has undergone severe eutrophication followed by a green tide episode; two species of Ulva, rotundata Blid. and Ulva curvata (Kütz.) De Toni, were the main macroalgae responsible for this bloom. From November 1993 to December 1994, we followed the biomass, the growth dynamics, and tissue elemental composition (C:N:P)of Ulva species, as well as some physicochemical variables in the estuary. Maximum biomass (up to 375 g dry wt·m^?2 in some spots, corresponding to a thallus area index of nearly 17 m²Ulva·m^?2 sediment) were observed in June and December. However, the biomass varied among the sampling stations. Water nitrate, ammonia, and phosphate showed high concentrations throughout the year, with extremely high transient pulses, sustaining the high growth rates observed. Growth rates were estimated directly in the field. The rates were generally higher in Ulva discs maintained in net cages than those estimated by changes in biomass standing stock between two consecutive samplings. The difference between both estimates was used to quantify the importance of the processes causing loss of biomass, which were attributable to grazing, exported biomass, and thallus decomposition under anaerobic conditions resulting from extreme self-shading. Maximum chlorophyll content was found in winter, whereas the minimum was in spring. Atomic N:P ratios were generally higher in the algae than in the water. However, the absolute concentrations of tissue N and P were always higher than the critical levels for maximum growth, which suggests that growth was not limited by inorganic N or P availability. The results suggested that the increase in nutrient loading in the river may have triggered the massive development of green algae and that light limitation and temperature stress in summer seem to be the main factors controlling the abundance of Ulva in the estuary. In addition to light availability and thermal stress, the different loss processes may have a decisive role in the dynamics of Ulva biomass.     

Improved isolation of glucuronan from algae and the production of glucuronic acid oligosaccharides using a glucuronan lyase

A new source for the production of bioactive glucuronic acid oligosaccharides (GlcUAOs) from the depolymerization of green seaweed Ulva lactuca glucuronan (Algal glucuronan) has been investigated. Algal glucuronan purification was optimized by the acidic precipitation method which allowed us to separate the polysaccharide mixture extracted from the cell wall of Ulva lactuca using hot water containing sodium oxalate. A series of the GlcUAOs were obtained by enzyme degradation of algal glucuronan with a glucuronan lyase (GL) isolated from Trichoderma strain. The putative bioactive GlcUAOs generated were then purified by size-exclusion chromatography in gram quantity and characterized by ¹H/¹³C NMR spectroscopy and ESI-Q/TOF-mass spectrometry.     

Management of natural <Emphasis Type="Italic">Ulva</Emphasis> spp. blooms in San Quintin Bay,Baja California: Is it justified?

According to Zertuche-González et al. (2009), Ulva spp. blooms, favored by oyster cultivation, are likely displacing subtidal meadows of Zostera marina in San Quintin Bay, Baja California. The authors propose a partial removal of the seaweed to reduce the risk of eutrophication and eelgrass displacement in the bay. We warn about the removal of Ulva spp. biomass by raising six arguments that emphasize the necessity of a historical and ecosystem-based management for San Quintin Bay. First, processes other than competitive exclusion by Ulva spp. blooms more likely explain changes of Z. marina subtidal meadows in the past decade. Second, there is no consistent evidence that oyster cultivation is promoting blooms of Ulva spp. and the loss of eelgrass. Third, the removal of Ulva spp. biomass is based on experiences of heavily anthropogenically eutrophic systems, while San Quintin Bay is not. Fourth, the proposed course of actions to restore eelgrass meadows ignores general historical baselines of estuarine and coastal systems by confusing what it means to be “pristine.” Fifth, despite the important experimental evidence indicating strong top–down control in temperate seagrass meadows, Zertuche-González et al. (2009) underestimated the capacity of consumers in structuring dynamics of vegetated soft-bottom communities in San Quintin Bay. Sixth, Ulva expansa may exert positive effects on seagrass ecosystem properties and functions. Instead, we propose protection against the propagation of unsustainable practices in the bay, and the reintroduction of large consumers that are now absent in this ecosystem. An ecosystem-based analysis of the role of Ulva spp. on eelgrass dynamics is needed.     

Application of <Emphasis Type="Italic">Gracilaria lichenoides</Emphasis> (Rhodophyta) for alleviating excess nutrients in aquaculture

A study was conducted in Xiangshan Bay, Ningbo, China, using red alga Gracilaria lichenoides to alleviate nutrient pollution in shrimp (Litopenaeus vannamei) and fish (Epinephelus awoara) culture ponds. Our results showed that G. lichenoides was efficient at absorbing inorganic nitrogen (IN) and inorganic phosphate (IP), and maintained a more stable dissolved oxygen (DO) level. A total of 506.5 kg (1,013 kg ha⁻¹) of shrimp and 210.5 kg (421 kg ha⁻¹) of fish were harvested from the shrimp/algae (SA) and fish/algae (FA) ponds, respectively. Only 53.5 kg shrimp were harvested from the shrimp pond without Gracilaria (S) due to anoxic asphyxia, and 163 kg fish were harvested from the fish culture pond without Gracilaria (F). Compared with using microalgae, bioremediation by macroalgae has no risk of harmful algal blooms (HABs), and it is easy to control seaweed biomass. During the experiment, there was a better environmental condition (lower chemical oxygen demand, IN, IP and chlorophyll a concentrations) in the ponds with Gracilaria. Furthermore, Gracilaria spp. can be used as food for abalone or other aquacultured animals and thus enhance economic return.     

Current state of seaweed resources in Spain

Commercial seaweeds in Spain are harvested on the north and northwest coasts. They are mainly agarophytes and carrageenophytes (Gelidium spp. and some Irish moss-like species, respectively), although some Phaeophyceae species (Fucus spp. and Laminaria spp.) are also exploited for alginates. No industrial seaweed cultivation is carried out in Spain at present. Spain's total commercial seaweed harvest can be estimated at 6,528 ± 2,076 t dry wt year^–1. Gelidium spp. are by far the most harvested, attaining 5,135 ± 1,761 t dry wt y^–1. To date, Spain's commercial seaweed harvest is low compared with that of other countries, but data on field standing crops and productivities of commercial taxa suggest that harvesting could be increased greatly.