Phycological research in the development of the Chinese seaweed industry

The term seaweed industry is employed in a broad sense and includes production both of commercial seaweeds such as Laminaria and Porphyra by phycoculture and of processed seaweed products, such as algin, agar and carrageenan.Before the founding of the People's Republic, China had a very insignificant seaweed industry, producing small quantities of the purple laver Porphyra and the glueweed Gloiopeltis by the primitive rock-cleaning method and the kelps Laminaria and Undaria by the primitive stone-throwing method, both aiming at enhancing the growth of the wild seaweeds. Also, a small quantity of agar was manufactured by the traditional Japanese method of gelling, freezing, thawing and drying the product. The small production was not sufficient to meet the demand of the Chinese people who for ages have appreciated seaweeds and their products for food. Therefore, large quantities of seaweeds and seaweed products had to be imported from various countries, for instance, Eucheuma and Gracilaria from Indonesia and other southeastern Asian countries, Laminaria and agar from Japan, even Porphyra from the USA. Annual Laminaria import from Japan generally amounted to over 10 000 tons and in some years approached 20 000–30 000 tons. Some quantities of the glueweed Gloiopeltis and the vermifuge weed Digenea simplex were exported, mainly to Japan.Since the founding of the People's Republic of China in October, 1949, China has exerted efforts to build up a self-supporting seaweed industry. Now after a lapse of 30-some years, a sizable seaweed industry has been developed. China is now able to produce by phycoculture more than one million tons of fresh seaweeds, including Laminaria, Undaria, Porphyra, Eucheuma, Gracilaria etc. and several thousand tons of seaweed extracts, including algin, agar, carrageenan, mannitol and iodine. At present, China still imports some quantities of seaweeds and seaweed products from various countries but is able to produce sufficient quantities to meet the people's need and even to export some quantities of the seaweeds Laminaria, Undaria and Porphyra and the seaweed products algin and mannitol.At the Tenth International Seaweed Symposium, I presented a paper on the Marine Phycoculture of China, in which I emphasized on the methods of cultivation (Tseng 1981b). Therefore I would like to take this opportunity to supplement the last lecture by presenting a paper on the role of phycological research in the development of China's seaweed industry.     

A Multinucleated Marine Amoeba Which Digests Seaweeds1

Marine amoebae were isolated during a search for organisms which degrade cell walls of seaweed. One of the isolates, a multinucleated amoeba (referred to here as Amoeba-I-7 or Am-I-7) was isolated from live tissues of the brown seaweed Sargassum muticum. It digested a variety of brown and red seaweeds including their walls and cuticles. Axenic clone cultures were isolated from cells that migrated on agar. Cultures were grown on agar or in liquid media. Seaweeds, seaweed wall extracts, and unicellular algae were tested as food sources.     

Seaweed Oligosaccharides Stimulate Plant Growth by Enhancing Carbon and Nitrogen Assimilation, Basal Metabolism, and Cell Division

It is now well established that plant cell wall oligosaccharides can stimulate or inhibit growth and development in plants. In addition, it has been determined that seaweed (marine algae) cell wall polysaccharides and derived oligosaccharides can enhance growth in plants. In particular, oligo-alginates obtained by depolymerization of alginates from brown seaweeds increase growth of different plants by enhancing nitrogen assimilation and basal metabolism. Interestingly, oligo-alginates also stimulate growth of marine and fresh water green microalgae, increasing the content of fatty acids. On the other hand, oligo-carrageenans obtained by depolymerization of carrageenans from red seaweeds increase growth of tobacco plants by enhancing photosynthesis, nitrogen assimilation, basal metabolism, and cell division. In addition, oligo-carrageenans increase protection against viral, fungal, and bacterial infections in tobacco plants, which is determined, at least in part, by the accumulation of several phenylpropanoid compounds (PPCs) with antimicrobial activity. Moreover, oligo-carrageenans stimulate growth of 3-year-old Eucalyptus globulus trees by increasing photosynthesis, nitrogen assimilation, and basal metabolism. Furthermore, oligo-carrageenans induce an increase in cellulose content and in the level of essential oil and some PPCs with antimicrobial activities, suggesting that defense against pathogens may be also enhanced. Thus, seaweed oligosaccharides induce a dual beneficial effect in plants and trees, enhancing growth, which is determined by the increase in carbon and nitrogen assimilation, basal metabolism, and cell division, and defense against pathogens, which is determined by the accumulation of compounds with antimicrobial activities. In this sense, molecular mechanisms that potentially interconnect activation of plant growth and defense responses are discussed.     

Cultivation and utilisation of red seaweeds in the Western Indian Ocean (WIO) Region

Seaweed farming in the Western Indian Ocean (WIO) Region is carried out in a number of countries, most of them farming Eucheuma denticulatum, Kappaphycus alvarezii and Kappaphycus striatum. These species are farmed mostly in Tanzania with limited production in Madagascar, Mozambique and Kenya; current production (2012) stands at 15,966 t (dry weight) year^?1 of Eucheuma and Kappaphycus, valued at US$ 4.2 million with 95 % of this tonnage coming from Tanzania. Other countries in the region have limited or no seaweed production owing to problems of epiphytes, ice ice and markets. The problem of epiphytes coupled with ice ice that WIO countries are facing causes die-off of Kappaphycus which is the preferred species in foreign markets for its thicker gel, kappa carrageenan (vs. the weaker iota carrageenan from Eucheuma). New efforts are put to curb these problems including moving seaweed farms to deeper waters and cultivation trials of other carrageenophytes as well as agar-producing species, agarophytes. Research work has been initiated to evaluate Gracilaria and Hypnea farming and processing in Tanzania, the Republic of Mauritius and Mayotte. Gracilaria farming is at experimental stages as a biofilter of fishpond effluents and as potential species for the production of agar with growth rates of 1.5–1.9 % day^?1. Hypnea farming is only being initiated in Mauritius and Mayotte at present. Other innovations including value addition by making various seaweed products and encouraging the consumption of seaweed as food at least in Tanzania and Mauritius are increasing further the importance of the seaweed farming and processing industry in the WIO Region.     

The seaweed hydrocolloid industry: 2016 updates,requirements, and outlook

The seaweed hydrocolloid industry, comprising agar, alginate, and carrageenan extracts, continues to grow in the order of 2–3% per year with the Asia-Pacific region increasingly dominating the raw material and manufacturing aspects of the industry. Geographic overviews, also in a historical perspective, of seaweed raw material availability including prices and consumption, manufacturing capacities, and utilizations and sales of extracts is presented. Some current and future industry dynamics, requirements, and changing structures, e.g., Indonesia’s increasingly dominant role within farming of agar and carrageenan-bearing seaweed species, randomly imposing of seaweed harvest restrictions or ban on exports, creation of a global certification standard for seaweed, and supply-demand dynamics for seaweed versus future global population are presented. The industry is increasingly being commoditized and China has become an important and, in many cases, dominant factor within all types of seaweed hydrocolloids and some explanations to this and strategic response by the rest of the industry is also touched upon. Also presented are some areas where the seaweed industry needs help from the scientific community. The main challenge is the ongoing general seaweed deterioration experienced in cultivated species—how are the strains to be improved and revitalized and can cultivation techniques be improved further? There is a general trend towards sustainability and, although seaweed cultivation and harvest can be sustainable, there is interest in the development of greener processes.     

Seaweeds of potential economic importance in Kenya: field survey and future prospects

Kenya is a net importer of agar and alginate based on recent government statistics, although it may have the potential to be self-sufficient or even an exporter of these phycocolloids. There is little information on carrageenan importation into Kenya since government statistics incorporate it as agar. Seaweeds are relatively unimportant in the Kenyan diet since they are consumed rarely by coastal people. A survey of 15 sites along of the Kenyan coast evaluated the potential for harvesting seaweeds and for establishing seaweed farms. Gracilaria appears to comprise the bulk of the low grade agar import, even though local species of this genus are widely distributed along the Kenyan coast. Major populations of Gelidium may be a potential source of high quality bacteriological grade agar. Eucheuma may be farmed locally to support the increasing local and regional demands for carrageenan. Recommendations for a national program of management and production for Kenya will be discussed.     

Worldwide distribution of commercial resources of seaweeds including Gelidium

The commercial exploitation of seaweeds for use as food and for the production of agar, alginate and carrageenan is outlined. The quantities of seaweed harvested for each purpose are tabulated and discussed. Seaweeds for food are derived chiefly from China, Japan and Korea, with almost 94% obtained by cultivation. Alginophytes are collected in 15 countries but six of these account for more than 80% of the total harvest; all are from natural stocks except for a large quantity of Laminaria cultivated in China. Natural carrageenophytes, from 12 countries, now account for only 20% of the total harvest; the remainder is cultivated Eucheuma species, 99% of which is produced in only two countries, the Philippines and Indonesia. Of the four categories of commercial resources of seaweeds considered, agarophytes are spread more evenly over a greater number of countries; they come from 20 countries and only five of these are minor contributors to the total. Gelidium species are particularly important because of the high quality agar they yield; their distribution and location are discussed.     

Floating seaweed and the influences of temperature, grazing and clump size on raft longevity — A microcosm study

Laboratory experiments were conducted to evaluate the longevity, and consequently also the rafting capacity of the brown seaweeds Fucus vesiculosus and Ascophyllum nodosum. The seaweed degradation process and the activity of the grazer Idotea baltica were strongly influenced by temperature: only at 5 °C, the seaweed growth exceeded the weight loss. At higher temperatures, seaweed fragments sank quickly (within 100 days at temperatures higher than 15 °C). This process was significantly accelerated in the presence of I. baltica, resulting in a decrease of raft longevity of 60-70%. At a constant temperature of 15 °C and in the absence of grazers, fragments of A. nodosum floated longer (mean 45 weeks) than fragments of F. vesiculosus (mean 15 weeks). The results indicate that floating seaweeds have the potential to stay afloat for a long time, but that their longevity is temperature-dependent and can be strongly reduced by grazing activity of associated herbivores.     

Mesoherbivores reduce net growth and induce chemical resistance in natural seaweed populations

Herbivory on marine macroalgae (seaweeds) in temperate areas is often dominated by relatively small gastropods and crustaceans (mesoherbivores). The effects of these herbivores on the performance of adult seaweeds have so far been almost exclusively investigated under artificial laboratory conditions. Furthermore, several recent laboratory studies with mesoherbivores indicate that inducible chemical resistance may be as common in seaweeds as in vascular plants. However, in order to further explore and test the possible ecological significance of induced chemical resistance in temperate seaweeds, data are needed that address this issue in natural populations. We investigated the effect of grazing by littorinid herbivorous snails (Littorina spp.) on the individual net growth of the brown seaweed Ascophyllum nodosum in natural field populations. Furthermore, the capacity for induced resistance in the seaweeds was assessed by removing herbivores and assaying for relaxation of defences. We found that ambient densities of gastropod herbivores significantly reduced net growth by 45% in natural field populations of A. nodosum. Seaweeds previously exposed to grazing in the field were less consumed by gastropod herbivores in feeding bioassays. Furthermore, the concentration of phlorotannins (polyphenolics), which have been shown to deter gastropod herbivores, was higher in the seaweeds that were exposed to gastropod herbivores in the field. This field study corroborates earlier laboratory experiments and demonstrates that it is important to make sure that the lack of experimental field data on marine mesoherbivory does not lead to rash conclusions about the lack of significant effects of these herbivores on seaweed performance. The results strongly suggest that gastropods exert a significant selection pressure on the evolution of defensive traits in the seaweeds, and that brown seaweeds can respond to attacks by natural densities of these herbivores through increased chemical resistance to further grazing.     

Volatile halocarbon emissions by three tropical brown seaweeds under different irradiances

The emission rates of eight volatile halogenated compounds by three tropical brown seaweed species collected from Cape Rachado, west coast Peninsular Malaysia, under different irradiances have been determined. A purge-and-trap sample preparation system with a gas chromatograph and mass-selective detector was used to measure a suite of halocarbons released by Sargassum binderi Sonder ex J. Agardh, Padina australis Hauck, and Turbinaria conoides (J. Agardh) Kützing. All species are widely distributed in Peninsular Malaysia, with S. binderi a dominant seaweed species at our survey site. Release of few halocarbons was found to be influenced by irradiance. Correlations were also observed between emission of certain halocarbons with photosynthetic activity, especially bromo-and iodinated compounds (0.6?<?r?<0.9; p?<?0.01) suggesting that environmental factors such as light can affect the release of these volatile halogenated compounds by the seaweeds into the atmosphere. Compared with temperate and polar brown seaweeds, tropical species, such as T. conoides, may emit higher levels of bromoform, CHBr₃, and other halocarbons. It is therefore important to investigate the contribution of tropical seaweeds towards the local atmospheric composition of halocarbons.     

Historical context of commercial exploitation of seaweeds in Brazil

Harvested by coastal populations for centuries, seaweeds have played an important role in the economy of a number of countries. In Brazil, they occur along the coastline, but are more diversified and abundant from the northeast to a portion of the southeast coast. Historically, the seaweed industry in Brazil is based on seaweed harvesting of natural beds. This practice continues to this day in a number of coastal communities in Northeastern Brazil. Since the 1960s, species of the genera Gracilaria and Hypnea have been collected in the intertidal zone for extraction of agar and carrageenan. Maximum production was achieved in 1973–1974, a period in which the country exported around 2000 t annually (dry weight) to Japan. Later (1977–1979), there was a sharp drop and annual exports fell to 250 t (dry weight). In 1981, Brazil exported only 150 t of dried seaweed for agar extraction. Between 1990 and 2000, overexploitation, decline in a number of agarophyte populations, poor quality, low price, and lack of a socioeconomic policy led to the almost total disappearance of this industry in Northeastern Brazil. Seaweed harvesting on natural beds is currently in decline, and the population that depended on this resource had to migrate or convert to other economic activities, such as fishing, aquaculture, and underwater tourism. However, the promising results obtained in pilot projects (Gracilaria and Kappaphycus) show that Brazil has significant potential as a seaweed biomass producer.     

Microbial alginate production,modification and its applications

Alginate is an important polysaccharide used widely in the food, textile, printing and pharmaceutical industries for its viscosifying, and gelling properties. All commercially produced alginates are isolated from farmed brown seaweeds. These algal alginates suffer from heterogeneity in composition and material properties. Here, we will discuss alginates produced by bacteria; the molecular mechanisms involved in their biosynthesis; and the potential to utilize these bacterially produced or modified alginates for high-value applications where defined material properties are required.     

Rapid determination of nucleosides,nucleobases and free amino acids in brown seaweeds using ultra-performance liquid chromatography coupled with triple quadrupole mass spectrometry

In this study the contents of nucleosides, nucleobases and free amino acids (FAAs) in common brown seaweeds Saccharina japonica, Sargassum pallidum, S. fusiforme, S. thunbergii and S. muticum from major production areas in China were determined by ultra-performance liquid chromatography coupled with triple quadrupole mass spectrometry (UPLC-TQ-MS). The total contents of nucleosides and nucleobases varied from 48.46 to 4,105.00 μg g^?1. Among the five seaweeds, levels of nucleosides and nucleobases in S. japonica were higher than the other species. The contents of ribonucleosides in these brown seaweeds were higher than those of deoxynucleosides. Uridine was the predominant constituent in all samples while 2′-deoxycytidine was the least in most samples. FAAs contents ranged from 0.14 to 22.76 mg g^?1. Among the five seaweeds, levels of FAAs in S. muticum were higher than the other species. Essential amino acids accounted for 3.26–51.49 % of total FAAs contents in different species. Glutamic acid was the major constituent in most samples. Four non-protein amino acids, γ-aminobutyric acid, taurine, hydroxyproline and citrulline, were found in the seaweed samples. Furthermore, principal component analysis (PCA) showed that chemical profiles of nucleosides, nucleobases and FAAs in S. pallidum and S. fusiforme were different from S. muticum, S. thunbergii and S. japonica. Common brown seaweeds could be promising natural sources for future industrial research into nucleosides, nucleobases and FAAs with potential benefits for human health.     

A novel high-performance anion-exchange chromatographic method for the analysis of carrageenans and agars containing 3,6-anhydrogalactose

A novel method has been developed to determine the sugar composition of 3,6-anhydrogalactose-containing polysaccharides, such as carrageenan and agar. The method is based on reductive hydrolysis with a methylmorpholine-borane complex in the presence of acid and subsequent high-performance anion-exchange chromatography analysis of the alditols without any derivatization. The method was validated by 13C NMR analysis of six carrageenans and three agars and by a previously used method based on derivatization to alditol acetates and gas-liquid chromatography analysis. The new method was found to be superior to the gas-liquid chromatography method as the analysis time was less than half. Also it was found to be more accurate and reproducible and no derivatization was required. The analysis of the six different carrageenan samples revealed that homogeneous mu- and nu-carrageenan, theoretically without 3,6-anhydrogalactose residues, cannot be isolated from red seaweeds. Consequently, the question arose if mu- and nu-carrageenans at all are present in seaweeds and if the current hypotheses regarding biosynthesis of carrageenans in the seaweeds are correct. The data demonstrated that carrageenans are highly complex natural polysaccharides, which are more irregular than assumed hitherto. The new analytical technique will permit elucidation of the detailed structure of seaweed polysaccharides and determination of their structure-property relationships.     

Discolored Red Seaweed <Emphasis Type="Italic">Pyropia yezoensis</Emphasis> with Low Commercial Value Is a Novel Resource for Production of Agar Polysaccharides

The red seaweed Pyropia yezoensis has been demonstrated to be a novel resource for the production of high-quality agar. P. yezoensis is grown for the food industry in large-scale Japanese mariculture operations. However, discolored P. yezoensis is mostly discarded as an industrial waste, although it has some kind of utility values. Here, we evaluated the utility of discolored P. yezoensis as a resource for agar production. The quality of agar from the discolored seaweed was comparable to that from normal seaweed. In addition, as a distinguishing characteristic, agar yield was higher from discolored seaweeds than from normal types. Moreover, we successfully used agar from discolored P. yezoensis for bacterial plate media and DNA electrophoresis gels without agarose purification. Thus, our results demonstrate that discolored P. yezoensis is suitable for agar production and use in life science research. Diverting discolored P. yezoensis from disposal to agar production provides a solution to the current industrial waste problem in mariculture, as well as a secure source of agar for research purposes.     

Bioethanol production from the hydrolysate of Palmaria palmata using sulfuric acid and fermentation with brewer’s yeast

Seaweeds, particularly species of red macroalgae, are promising resources for bioethanol production because of their exceptionally high carbohydrate content. Of 20 seaweeds evaluated, Palmaria palmata (Rhodymenia palmata) contained the highest carbohydrate content (469.8 mg g^?1 seaweed) with a carrageenan content of 354 mg g^?1 seaweed. Such a high carrageenan content makes the high-volume production of bioethanol feasible. Acid hydrolysis of P. palmata in 0.4 M H₂SO₄ at 125 °C for 25 min released 27 mg of glucose, 218.4 mg of reducing sugars, and 127.6 mg of galactose per gram of seaweed. Ethanol fermentation of these hydrolysis products using an inoculum concentration of 1.5 mg mL^?1 at 30 °C and 72 h in a shaking incubator at 130 rpm yielded 17.3 mg of ethanol per gram of seaweed.     

Stocks and the use of economic marine macrophytes of Vietnam

This review analyzes the literature and original data on the resources, use, and cultivation of useful marine macrophytes in Vietnam. Information is provided about more than 60 species of macroalgae: including 41 species of Rhodophyta, 15 species of Heterokontophyta and 7 species of Chlorophyta. Out of these, more than 50 species are used for food, about 20 species are used in folk medicine, and more than 30 species are important commercial sources of polysaccharides (agar, carrageenan, and alginates); Sargassum, Gracilaria, and Kappaphycus are exported (mainly to China). The wild stocks of useful algae in Vietnam are declining as a result of their uncontrolled export (especially Sargassaceae). Several species of Gracilaria and Kappaphycus are commercially cultivated and also exported.     

High-value products from macroalgae: the potential uses of the invasive brown seaweed<Emphasis Type="Italic">, Sargassum muticum</Emphasis>

Marine seaweeds represent an abundant source of natural products and may harbour valuable chemicals. The brown seaweed Sargassum muticum is an invasive species to the coasts of the British Isles, mainland Europe and North America. Attempts at its eradication and control have generally not been successful, although time-consuming and costly. Commercial exploration of this biomass for food, fuel and pharmaceutical products could encourage its harvesting and control. Though S. muticum might be unsuitable as a source of biofuel due to high ash and water content, this rapidly growing macroalga has a naturally high content of antioxidants, carotenoids and phenols, including the well-known anti-cancer compound fucoxanthin, making this species a potential source of a range of pharmaceutically relevant materials.     

Detection of Differential Host Susceptibility to the Marine Oomycete Pathogen Eurychasma dicksonii by Real-Time PCR: Not All Algae Are Equal

In the marine environment, a growing body of evidence points to parasites as key players in the control of population dynamics and overall ecosystem structure. However, their prevalence and impact on marine macroalgal communities remain virtually unknown. Indeed, infectious diseases of seaweeds are largely underdocumented, partly because of the expertise required to diagnose them with a microscope. Over the last few years, however, real-time quantitative PCR (qPCR) has emerged as a rapid and reliable alternative to visual symptom scoring for monitoring pathogens. Thus, we present here a qPCR assay suitable for the detection and quantification of the intracellular oomycete pathogen Eurychasma dicksonii in its ectocarpalean and laminarialean brown algal hosts. qPCR and microscopic observations made of laboratory-controlled cultures revealed that clonal brown algal strains exhibit different levels of resistance against Eurychasma, ranging from high susceptibility to complete absence of symptoms. This observation strongly argues for the existence of a genetic determinism for disease resistance in brown algae, which would have broad implications for the dynamics and genetic structure of natural populations. We also used qPCR for the rapid detection of Eurychasma in filamentous brown algae collected in Northern Europe and South America and found that the assay is specific, robust, and widely applicable to field samples. Hence, this study opens the perspective of combining large-scale disease monitoring in the field with laboratory-controlled experiments on the genome model seaweed Ectocarpus siliculosus to improve our understanding of brown algal diseases.     

Alginate stability during high salt preservation of Ascophyllum nodosum

Formaldehyde is usually added to brown algae to avoid microbial growth during storage and to fix polyphenols in the algae before alginate extraction. Since formaldehyde is toxic, allergenic and possibly carcinogenic, dry salting of Ascophyllum nodosum was tested as an alternative. The seaweeds, harvested at locations with a salinity of about 30‰ from late autumn to early spring, were stored at 22±2 °C under compost-like conditions. Untreated samples of seaweed lost their quality as a raw material for alginate production within 14 days. Salted (20–22%) as well as formaldehyde treated seaweed was preserved for at least 46 days. Due to the reduced water activity and oxygen saturation in the dry salted seaweed, microbial growth and brown colouring reactions were suppressed. Economic factors must also be taken into account before large-scale applications are considered. This revised version was published online in June 2006 with corrections to the Cover Date.