Seaweed production: overview of the global state of exploitation,farming and emerging research activity

ABSTRACT

The use of seaweeds has a long history, as does the cultivation of a select and relatively small group of species. This review presents several aspects of seaweed production, such as an update on the volumes of seaweeds produced globally by both extraction from natural beds and cultivation. We discuss uses, production trends and economic analysis. We also focus on what is viewed as the huge potential for growing industrial-scale volumes of seaweeds to provide sufficient, sustainable biomass to be processed into a multitude of products to benefit humankind. The biorefinery approach is proposed as a sustainable strategy to achieve this goal. There are many different technologies available to produce seaweed, but optimization and more efficient developments are still required. We conclude that there are some fundamental and very significant hurdles yet to overcome in order to achieve the potential contributions that seaweed cultivation may provide the world. There are critical aspects, such as improving the value of seaweed biomass, along with a proper consideration of the ecosystem services that seaweed farming can provide, e.g. a reduction in coastal nutrient loads. Additional considerations are environmental risks associated with climate change, pathogens, epibionts and grazers, as well as the preservation of the genetic diversity of cultivated seaweeds. Importantly, we provide an outline for future needs in the anticipation that phycologists around the world will rise to the challenge, such that the potential to be derived from seaweed biomass becomes a reality.     

Carbon nutrition of seaweeds: Photosynthesis,photorespiration and respiration

Techniques are described for measuring gas exchange in seaweeds held in moist air (air suspension). In the species we have examined, oxygen has little or no effect on photosynthesis except at very low (50 μ1·1^?1) CO₂ concentration. Photorespiration could not be detected unless the seaweeds were treated simultaneously with high O₂ and low CO₂ or with the carbonic anhydrase inhibitor, diamox. However, sporulating and meristematic tissues exhibit oxygen-insensitive light respiration (CO₂production in light not associated with photorespiratory metabolism). Elevated pH in the surface water of seaweeds also caused light respiration. Oxygen-sensitive wound respiration was observed that could easily be mistaken for photorespiration. C₄ photosynthesis could not be detected. On the basis of several experimental approaches it was concluded that these seaweeds normally absorb bicarbonate rather than CO₂ from sea water. High CO₂ concentrations are required in gas streams aerating seaweed cultures in air or water suspension to maintain the bicarbonate concentration at levels normally found in sea water and to support normal levels of photosynthesis.     

Floating period of Sargassacean thalli estimated by the change in density

The density (fresh weight/volume) of the detached thalli of four sargassacean species (Myagropsis myagroides, Sargassum horneri, S. patens, S. siliquastrum) was measured weekly to determine their floating period. Thalli of the four species were collected with respect to the difference in their reproductive stage, and their floating abilities at different stages were determined. Floating thallus was ballasted with pre-weighed leads until the final addition of a weight caused sinking. The density and floating period of sargassacean thalli were dependent on the species and the specimens’ reproductive stage. The floating period of thalli detached before maturation was estimated to be between 4 and 14 weeks. However, thalli detached during maturation had a floating period of between 1 and 8 weeks, and those detached after maturation sank within 2 weeks. Sargassum horneri had the lowest density and the longest floating period among the four species investigated and this may explain the high abundance of S. horneri in floating seaweeds observed in the Sea of Japan.     

An overview of seaweed resources in Chile

The seaweed harvest in Chile has doubled during the past decade, and export values have increased by 300% because of diversification and increase in the volume of products with greater value added. The export value of seaweed products increased from US $18 million in 1980 to $52 million in 1991. During the past decade, the successful cultivation of Gracilaria was implemented, and this has compensated for the large decrease in yields from natural beds. In the short term, it will be necessary to develop techniques for the cultivation of other resources such as Iridaea, Gigartina, Lessonia and Gelidium. Alternative biotechnological methods must also be developed, such as the use of Gracilaria strains with increased quality and production for growth in cultivation centers.     

Gelidium commercial exploitation: natural resources and cultivation

The state of the Gelidium agarophyte resource was last reviewed in 1990 during the First International Workshop on Gelidium. The main objective of the present study was to gather and analyse the new information made available since then, with emphasis on the G. sesquipedale resource from the Iberian Peninsula and Morocco. Other Gelidium-based resources world-wide are also reviewed, as well as new attempts to cultivate Gelidium species in the sea. In general, since the late 1980s, G. sesquipedale yields have decreased in Portugal, and increased in Spain and Morocco. Current reported values, in dry weights, are ca. 670 t yr-1 for Portugal (average 1995–1997), 5200 t yr-1 in Spain, and 6950 t yr-1 for Morocco (average 1994–1996). Efforts to develop cultivation of G. sesquipedale in the sea in Spain have yet to progress beyond the demonstration scale. Other important Gelidium-based agarophyte resources are located in: (i) South Africa where, in addition to G. pristoides, new species are being harvested, with total agarophyte landings of 140 t yr-1 in 1996; (ii) Mexico, where G. robustum harvest averaged 750 t yr-1 in 1987–1996; (iii) and Chile where harvest of Gelidium species yielded ca. 460 t yr-1 in 1990–1992. Preliminary research on new techniques for commercial cultivation of Gelidium species is also reported from these countries. This revised version was published online in June 2006 with corrections to the Cover Date.     

Sustainable harvesting of wild seaweed resources

Abstract

Macroalgae have played an important role in coastal communities for centuries. In the past, they have been harvested and gathered from shorelines around the world for traditional uses such as food, animal feed and a crude fertilizer (marine manure). Today, seaweeds are used in a multitude of applications with expanding global industries based on hydrocolloids, cosmetics and food supplements, and also as a potential biofuel source. However, of the approximately 10?000 algal species reported to exist, only a small number are commercially utilized. While representing only a small fraction of total global seaweed production, harvesting and gathering ‘wild’ seaweeds has had, and continues to have, an integral role in many coastal societies, often being intrinsically linked to the cultural identity of those coastal communities. Today, 32 countries actively harvest seaweeds from wild stocks, with over 800?000 t harvested annually from natural beds. It is vitally important that seaweeds are utilized sustainably and that natural resources are effectively managed by coastal communities with vested interests around the world. As the popularity of seaweeds increases and the use of less traditional species with novel applications comes to the fore, it is critically important to make certain that the sustainability of the resource is ensured given the increased pressures of harvesting. Issues exist regarding ownership of the resource and its over-exploitation, and the implementation of environmentally damaging harvesting techniques must be avoided. Resource scientists, managers, conservationists, governments, and other stakeholders need to be proactive in the sustainable management of these vulnerable, yet valuable, resources.     

THE PERFORMANCE OF SPRAY‐IRRIGATED ULVA LACTUCA (ULVOPHYCEAE,CHLOROPHYTA) AS A CROP AND AS A BIOFILTER OF FISHPOND EFFLUENTS1

The seaweed Ulva lactuca L. was spray cultured by mariculture effluents in a mattress‐like layer, held in air on slanted boards by plastic netting. Air‐agitated seaweed suspension tanks were the reference. Growth rate, yield, and ammonia‐N removal rate were 11.8% · d^?1, 171 g fresh weight (fwt) · m^?2 · d^?1, and 5 g N · m^?2 · d^?1, respectively, by the spray‐cultured U. lactuca, and 16.9% · d^?1, 283 g fwt · m^?2 · d^?1, and 7 g N · m^?2 · d^?1, respectively, by the tank U. lactuca. Biomass protein content was similar in both treatments. Dissolved oxygen in the fishpond effluent water was raised by >3 mg · L^?1 and pH by up to half a unit, upon passage through both culture systems. The data suggest that spray‐irrigation culture of U. lactuca in this simple green‐mattress‐like system supplies the seaweed all it needs to grow and biofilter at rates close to those in standard air‐agitated tank culture.     

Structural Features of N-Glycans of Seaweed Glycoproteins: Predominant Occurrence of High-Mannose Type N-Glycans in Marine Plants

An ice nucleation-active strain of Erwinia uredovora shed vesicles when cultured at low temperature (l0?C). We isolated an ice nucleation-active vesicle fraction from the culture medium by ultrafiltration, ultracentrifugation, and gel filtration. Western blot analysis showed that this cell-free vesicle fraction contained an ice nucleation-active protein (InaU). The process of the InaU transport to a shed vesicle was examined by immunohistochemical analysis using electron microscopy. The examination showed the following successive processes: InaU molecules first assemble around the inner membrane, then the assembly enters a vesicle justforming on the surface of the outer membrane, and finally the vesicle, 100–400 nm in diameter, leaves the surface to be shed with InaU molecules occluded.     

Seaweed research and utilization in Chile: moving into a new phase

Two phases have been distinguished classically in the history of Latin American phycological research: the explorer phase characterized by the taxonomic work of mainly European and North American scientists, and the diversification phase marked by the establishment of resident scientists in the area and the training of a new generation of phycologists in subjects other than taxonomy. Over the last 15 years, Chile has entered a third phase, characterized by a significant increase in scientific and economic activity centered around seaweeds. Seaweed cultivation has been commercialized; raw materials are now locally processed and economic returns have more than tripled. In addition, some groups of opportunistic seaweed gatherers have become farmers. Loosely correlated with the above developments has been a significant increase in the number of scientific and technological studies related to seaweeds, in the number of professional phycologists and in the specialization of the various groups. This study first describes these new developments and the conceptual advances achieved in farming and resource management. It also emphasizes some socio-economic differences with seaweed farming in other countries and explores the level of interaction between the local scientific and productive sectors in view of future developments.     

Modeling competition between Laurencia obtusa (Ceramiales,Rhodophyta) and Hypnea spinella (Gigartinales,Rhodophyta) at Cabo Frio Island,Rio de Janeiro,Brazil

The Lotka-Volterra competition model was used to represent the interaction between Laurencia obtusa and Hypnea spinella. A new model that considers effects of competition on algal carrying capacity is suggested. To test the models, data from field experiments conducted in an intertidal region at Cabo Frio Island, Rio de Janeiro, Brazil, were used. Both models showed that Hypnea was a stronger competitor than Laurencia. The model of interaction through the carrying capacity showed a stable coexistence between the algal populations and better represented the experimental data.     

Spatial and seasonal variations in the biofunctional lipid substances (fucoxanthin and fucosterol) of the laboratory-grown edible Japanese seaweed (Sargassum horneri Turner) cultured in the open sea

This work studied the effect of spatial and seasonal differences on the accumulation of functional lipid components in Sargassum horneri (Turner), an edible Japanese seaweed popularly called Akamoku. S. horneri obtained from Samenoura bay area of Japan was laboratory cultured to evaluate the effect of temperature on the accumulation of total lipids (TL), fucoxanthin (Fx) and fucosterol (Fs) by the alga. The laboratory cultured 3 month old S. horneri were cultured in the open sea in two different geographical locations off Usujiri and Matsushima to evaluate the monthly variations, over a year, in their TL, Fx and Fs contents. S. horneri grown off the Usujiri area accumulated the maximum TL close to 193 mg g⁻¹ dry weight during the coldest part of the year. Fx and Fs contributed 5.6% and 16.2% of the TL in S. horneri harvested off Usujiri in February. Further, in spite of being the same species and parent stock, S. horneri grown off the Matsushima area accumulated less TL, Fx and Fs as compared to their Usujiri counterparts. Our study clearly indicates the role of temperature and light apart from nutritional profile and depth of waters where the seaweed was grown on the accumulation of functional lipid components in S. horneri.     

Dynamics of antibacterial activity in three species of Caribbean marine algae as a function of habitat and life history

Antibacterial activity of lipid extracts from three species of Caribbean marine algae, Spyridia filamentosa, S. hypnoides and Wrangelia bicuspidata was evaluated monthly for one year. Activities were tested for whole plant extracts and TLC-separable zones. Whole plant extracts demonstrated monthly variability in activity with respect to both habitat and life history phase in addition to periods of similarity. No consistency was seen regarding activity against different microorganisms. TLC analyses of the extracts led to the identification of twenty-seven chromatographically distinct regions (TLC zones) each from both S. filamentosa and S. hypnoides and twenty-five from W. bicuspidata, which demonstrated antimicrobial activity. Between these species, twenty-one active TLC zones appeared to be shared based on their similar chromatographic characteristics. Individual TLC zones also demonstrated variable activity throughout the sampling period with respect to habitat and life history phase as well as periods of similarity. Algal antibiosis in these species is recognized as being highly complex, involving numerous chemical compounds, each of which is highly variable in terms of its presence and/or probable concentration.     

Structural study of fucoidan from Cladosiphon okamuranus tokida

A structural study was carried out on a fucoidan isolated from the brown seaweed Cladosiphon okamuranus. The polysaccharide contained fucose, glucuronic acid and sulfate in a molar ratio of about 6.1 : 1.0 : 2.9. The results of Smith degradation showed that this polysaccharide has a linear backbone of 13-linked -fucopyranose with a half sulfate substitution at the 4-positions, and a portion of the fucose residues was O-acetylated. The data obtained from partial acid hydrolysis, a methylation analysis and NMR spectra indicated that the -glucuronic acid residue is linked to the 2-positions of the fucose residues, which were not substituted by a sulfate group. These results indicated that the average structure of this fucoidan is as follows: -[(3Fuc-4(±OSO₃-)1–)₅3[GlcA12]Fuc1–]_n–. (Half of each fucose residue was sulfated. One O-acetyl ester was present in every 6 fucose residues.)     

Effect of seaweed concentrate on hydroponically grown spring barley

Spring barley (Hordeum vulgare cv. Triumph) was grown hydroponically over a 6-week period. Two treatments were incorporated either into the hydroponic solution or sprayed onto the plants at rates of 1 ml per 3 litres. The treatments applied were: (i) a seaweed concentrate prepared fromAscophyllum nodosum (L.) Le Jolis (marketed as Maxicrop Triple), (ii) a ‘Trace element’ treatment incorporating the micro and macro nutrients added to the seaweed extract base to produce the formulated product Maxicrop Triple and (iii) a control treatment. Irrespective of the mode of application, plants treated with Maxicrop Triple grew faster than plants under either of the two other treatments. Elevated growth rates were also found for the ‘Trace element’ treated plants when incorporated into the hydroponic solution. At the final harvest, plants with Maxicrop Triple incorporated into the hydroponic solution showed increases from 56–63% over the control treatment for the growth characteristics measured. ‘Trace element’-treated plants produced increases of between 25–45%. When the treatments were sprayed the effect was less pronounced. Maxicrop Triple increased growth characters by 35–38% and the ‘trace element’ treatment gave increases in the range of 2–13%.     

Fusion and histocompatibility in Rhodophyta

Fully developed thalli of Chondrus crispus, Gracilaria chilensis, Gymnogongrus furcellatus and Mazzaella laminarioides were used to assess tissue compatibility. The effect of thallus polarity on grafting and regeneration was also evaluated. Fusion did occur between fragments of the same life history phase in C. crispus, G. chilensis, G. furcellatus and M. laminarioides. Fusion between sporophytic and gametophytic tissue occurred in C. crispus, G. chilensis and M. laminarioides. Intergeneric fusion was observed between C. crispus and M. laminarioides, but not between G. chilensis and G. furcellatus.Outer cell wall, cortex and medulla were continuous at the contact face in compatible combinations. Medullary cells in the attached fragments were thinner and longer than normal cells, forming an interwoven scar plate. Thallus polarity did not modify fusion and regeneration.     

Effect of seaweed concentrate on the growth and mineral nutrition of nutrient-stressed lettuce

The effects of the seaweed concentrate Kelpak on the growth and mineral nutrition of lettuce plants grown under conditions of varying nutrient supply were investigated. Kelpak significantly increased the yield and the concentration and amounts of Ca, K and Mg in the leaves of lettuce receiving an adequate supply of nutrients, but had little effect on nutrient stressed plants. Results are discussed in relation to the physiological mechanisms that have been proposed to explain the beneficial effects of seaweed concentrate on plants.author for correspondence