Interaction mechanisms between Gracilaria chilensis (Rhodophyta) and epiphytes

Epiphytes can have a considerable effect on Gracilaria production, and Ulva is one of the commonest algal species identified as an epiphyte, reaching loads of 60% (g of epiphytes per g of Gracilaria) in the intertidal cultures of southern Chile. This study evaluates the relative importance of light reduction, addition of weight to the host thalli and nutrient depletion, as mechanisms determining the interaction effects of Ulva epiphytes on Gracilaria cultivation. Using field experiments undertaken during the main Gracilaria growth season (spring), we evaluate the mechanisms of epiphyte-host algae interaction by manipulating artificial epiphytes. The results indicate that Ulva can significantly depress Gracilaria biomass production and that the addition of weight to the host algae and the consequent dislodgement increase, appear to be the main mechanisms involved in the Ulva-Gracilaria interaction. However, the light reduction caused by the epiphytes can also partially explain the reduction in Gracilaria production. Nutrients depletion would not appear to fully account for the detrimental effects of Ulva over Gracilaria in intertidal farming areas of southern Chile.     

Cultivation of Gracilaria on the sea-bottom in southern Chile: a review

This review contains information about the cultivation techniques, strategies, problems and new challenges faced as well as an economic analysis of the income-producing capacity of Gracilaria farming, considering the variability of environmental systems where this alga is cultivated in southern Chile. The development of Gracilaria farming in Chile was made possible by an increased market demand, as well as the existence of basic knowledge that permitted the management of wild stocks and the initiation of cultivation practices. Subtidal cultivation systems appear to be more productive than intertidal systems and are less susceptible to wave action than intertidal cultivation areas. In relation to farming practices, this difference implies that planting and harvesting methods and strategies vary between habitats where cultivation is being carried out on a commercial scale. Several problems such as the environmental impact of different cultivation methods adopted by the farmers, the management of contaminating organisms and strain selection appear to be important and new areas for future research. Finally, an analysis of the income-producing capacity indicates that environmental differences also have important consequences for the management strategies of Gracilaria cultivation.     

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.     

Recent trends of seaweed production in Chile

In the last fourteen years the production of seaweeds in Chile has ranged from 74 000 to 229 000 wet metric tons per year and has included about twenty species belonging to Phaeophyta and Rhodophyta. The only source of this production has been the exploitation of natural beds, except for Gracilaria, which is the only case of commercial cultivation and contributes significant quantities to total production. Initially most of the raw material was exported but currently important quantities of Gracilaria and several carrageenophytes are being processed by local industry. Changes in production of the main resources are analyzed with consideration of potential demand, level of knowledge about natural beds, and the situation of total Gracilaria farming, in order to attempt predictions for the supply. Current possibilities of applying new technologies to cultivate other economically important Chilean seaweeds are also analyzed and discussed.     

Effects of stock loading and planting distance on the growth and production ofGracilaria chilensis in rope culture

Through experimental manipulation of stock loading and planting distances in rope culture, this study evaluates the hypothesis that optimum density for growth and production ofGracilaria chilensis vary within a bed. Small packages ofGracilaria disposed at short distances exhibit higher growth and production rates in the landward ends of ropes than larger packages of seaweed disposed at longer distances. In these experiments, the landward end of ropes faces estuarine conditions, with reduced salinities and increased sediment loads during low tides. The interacting effects of stocks loading and position in the rope can affect growth and production rates so much as to nullify the 3.5-fold summer increments in production that occur in these estuarine beds of southern Chile. It is concluded that through manipulation of planting distances and stock loading, the production of this species could be increased up to 35% in some areas.     

Seasonal Occurrences of Epiphytic Algae on the Commercially Cultivated Red Alga Kappaphycus Alvarezii (Solieriaceae, Gigartinales, Rhodophyta)

Common problems faced in farming of the red algal genus Kappaphycus/Eucheuma are “ice-ice disease” and the occurrence of epiphytes. Considerable work has been documented on “ice-ice disease” and it's mode of infection but limited information is available on the emergence of epiphytes. The present study addresses the phenomenon of epiphyte infection, its prevalence in commercially cultivated red alga, Kappaphycus alvarezii, and their variability associated with seasonality. Cultured seaweed became susceptible to epiphytes in the dry seasons (1) between March – June and (2) September – November. Findings revealed Neosiphonia savatieri (Hariot) M. S. Kim et I. K. Lee, as the dominant infecting epiphyte, representing up to 80–85% of the epiphyte present during peak seasons. Besides N. savatieri, Neosiphonia apiculata, Ceramium sp., Acanthophora sp. and Centroceras sp. were observed in smaller quantities. SEM (Scanning Electron Microscope) micrographs revealed the epiphyte's attachment to the host. Further histological study showed the extent of penetration of epiphytes into the host's cortex tissues and condition of its surrounding tissues. The outbreak of epiphytic filamentous red algae also correlated with drastic changes in seawater temperature and salinity during March– June and September – November.     

World-wide use and importance of Gracilaria

The world's first source of agar, from the middle of the seventeenth century, was Gelidium from Japan, but by the beginning of the twentieth century demand for the phycocolloid exceeded of the supply of this alga. Since then Gracilaria has played an important role in the production of agar. Currently agars are obtained from five genera in three orders of red algae and marketed as ‘natural agar’ in squares or strips or as ‘industrial agar’ in powder form. The development of production processes through alkaline hydrolysis of sulphates has allowed a good quality food agar to be obtained from Gracilaria. This does not show the synergistic reaction with locust bean gum apparent with Gelidium agar. The term ‘agaroids’ is applied to Gracilaria agars produced without alkaline hydrolysis of sulphates, with greater sulphate content and much less gel strength. Unlike Gelidium, Gracilaria has to be processed in a short period of time and cannot be allowed to remain in storage for use during years of lower availability. Statistics of imports of agarophytes to Japan during the last 10 years give an indication of the state of the market. During this period there was a marked reduction in Gracilaria imports, mainly from Chile, but also the Philippines, Indonesia and South Africa, mainly due to the overall increase in the capacity of agar production in Gracilaria-producing countries.     

Harvesting of the Kelp Ecklonia Maxima in South Africa Affects Its Three Obligate, Red Algal Epiphytes

In South Africa, more than 7000 t (f wt) of kelp (Ecklonia maxima) fronds are harvested annually to feed cultured abalone. Carpoblepharis flaccida, Gelidium vittatum and Polysiphonia virgata are conspicuous red algal epiphytes on older kelps and provide habitat and food for numerous animals. Over 4.5 y, we examined the effects of one destructive harvest of E. maxima on these 3 epiphytes. Two 20 × 20 m plots of kelp with similar epiphyte loads were demarcated. In one, all E. maxima sporophytes with stipes longer than 50 cm were harvested. The other plot served as a control. After 2.5 y the biomass of E. maxima in the harvested plot had recovered to control levels, but the epiphyte load (g epiphytes. Kg kelp⁻¹) was statistically lower in the harvested plot after 2.5 and 3.5 y, and only recovered after 4.5 y. While most commercial harvesters cut through the “heads” (primary blades) of the kelp, effectively killing them, a new, non-lethal method removes secondary blades 20–30 cm from their bases, leaving the meristems and primary blades intact. At 5 sites studied, G. vittatum and P. virgata were found almost entirely on stipes and primary blades, and harvesting only distal parts of secondary blades limited losses to about 50% of C. flaccida biomass. To protect epiphytes, non-lethal harvesting is recommended and permanent non-harvest zones have been established in addition to limiting kelp yields and disallowing harvesting in Marine Protected Areas.     

Ecological differences among Chilean populations of commercialGracilaria

Field farming ofGracilaria is gradually replacing use of the wild crop in Chile. The most popular planting method consists of establishing underground thallus systems for patches ofGracilaria on wave-sheltered, soft-bottom habitats. Commercial cultivation often involves transplanting vegetative material to distant places along the coast. This study shows that native populations possess some ecological differences that might affect cultivation success. Seven environmental factors were tested for their effects on growth of four geographically separated populations belonging to two species ofGracilaria. No specific or population differences were found under various temperature or salinity regimes. Specific differences occurred in responses to irradiance, daylength, sand burial and sand abrasion. Responses to epiphytes differed among both population and species. Epiphyte recruitment was inhibited by some populations and stimulated by others. Given the farming methods presently used in Chile, these results have ecological and economical importance, as they suggest that transplanting randomly from one to other type of habitat does not assure cultivation success.     

Recent status of Gracilaria cultivation in Taiwan

The recent decrease in Gracilaria culture production and value in Taiwan were evaluated from statistical data and from interviews with local fishermen. Reasons are: 1) during 1986–87, many Gracilaria culture ponds were transformed to grow grass shrimp (Penaeus monodon) in monoculture, but disease of the shrimp occurring soon after stopped such production and Gracilaria culture took over, but 2) due to manpower shortage, Gracilaria-farmers prefer to sell their crops to abalone farmers and not to agar factories. Since Gracilaria as abalone feed is cheeper than for agar production, the value of algal crop decreased.     

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.     

Opportunities and challenges for the development of an integrated seaweed-based aquaculture activity in Chile: determining the physiological capabilities of <Emphasis Type="Italic">Macrocystis</Emphasis> and <Emphasis Type="Italic">Gracilaria</Emphasis> as biofilters

Seaweed production is a reality in Chile. More than ten species are commercially used to produce phycocolloids, fertilizers, plant growth control products, human food or animal fodder and feed additives. These multiple uses of algae offer a number of possibilities for coupling this activity to salmon, abalone and filter-feeder farming. In this context, different experiments carried out in Chile have demonstrated that Gracilaria chilensis and Macrocystis pyrifera have great potential in the development of an integrated aquaculture strategy. The present Integrated Multi-Trophic Aquaculture (IMTA) approach study showed that Gracilaria can be cultured best at 1 m depth whereas Macrocystis has an especially good growth response at 3 m depth. Both species use available nitrogen efficiently. On the other hand, high intensities of solar radiation (UV and PAR) can be critical at low depths of cultivation, and our results indicate that both species show photosynthetic susceptibility mainly at noon during the summer. The demand of Macrocystis for abalone feeding is increasing, thus improving the opportunity for developing an integrated nutrient waste recycling activity in Chile. Although Gracilaria shows a higher nitrogen uptake capacity than Macrocystis, its market value does not yet allow a massive commercial scaling.     

Phenology of Gigartina skottsbergii (Gigartinaceae,Rhodophyta) in Ancud Bay,southern Chile

This study reports the variations in biomass and reproductive phenology in a natural bed of Gigartina skottsbergii over a period of a year and correlates these variations with the abiotic factors solar radiation, number of daylight hours, water temperature and salinity. The results obtained show an annual production cycle with maximum biomass values in spring-summer, correlated with high solar radiation and to a lesser extent with salinity. Sexual reproduction was maximum in autumn-winter, correlated with low temperature and short-day conditions. The population showed a large gametophytic dominance according to size and biomass, which suggests that there is not ecological equivalence between the life history phases of the species. Finally, based on the results, a restriction of harvesting to spring-summer is suggested.     

Using gigartinine to track the distribution of an alien species of <Emphasis Type="Italic">Gracilaria</Emphasis> in New Zealand

The presence of gigartinine has been used previously as a taxonomic marker for the positive identification in Manukau Harbour (west coast, Auckland) of Gracilaria sp., which has apparently been introduced to New Zealand waters and is easily confused morphologically with the native species, G. chilensis. Thirty two estuarine/harbour populations of Gracilaria in New Zealand were screened for the presence of gigartinine to further test the utility of gigartinine as a reliable species marker. DNA fingerprinting was used to confirm the identity of a subset of the specimens surveyed. Using genetic rather than chemical characterisation, it was discovered that Gracilaria sp. is also present in Orakei Basin (east coast, Auckland). Although a sample from the wild did not have the anticipated gigartinine content, tank cultivation of thalli from this population in an artificially elevated nitrogen environment allowed the plant to accumulate nitrogen as gigartinine. This confirmed the unusual ability of this species of Gracilaria to store nitrogen in this form, unlike the widespread, morphologically similar, G. chilensis.     

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.     

The effect of localised eutrophication on competition between Ulva lactuca (Ulvaceae,Chlorophyta) and a commercial resource of Gracilaria verrucosa (Gracilariaceae,Rhodophyta)

In spring (August) 1993 a bloom of Ulva lactuca appeared for the first time in Saldanha Bay, South Africa, and persisted through summer. Ulva wash-ups contaminated the beach and part of the commercial Gracilaria beach-cast had to be discarded. The biomass and distribution of Gracilaria and Ulva are described in relation to the seasonal water chemistry of the bay. Gracilaria survives in deeper water in summer by the pulsing of nutrients on an approximately 6-day cycle of movement of the thermocline that separates nutrient-rich bottom water from warm oligotrophic surface water. Ammonium-rich fish-factory discharge into this surface layer in a sector of the bay provided localised conditions for Ulva to out-compete Gracilaria at depths of 2–5m, demonstrating the powerful disruptive effect of eutrophication in this strongly stratified system.     

Cultivation of Gracilaria in outdoor tanks and ponds

This review deals with the major problems of unattached Gracilaria intensive cultivation in outdoor tanks and ponds. These problems are presented through the main variables affecting the Gracilaria annual yield and the updated solutions evolved. The physical variables include tank and pond structure, seawater characteristics such as velocity, agitation practice, exchange rate, and salinity, light characteristics such as quantity and quality, and temperature modelling. The chemical variables include nutrient composition and regime of application, and inorganic carbon supply with the pH changes involved. The biological variables include seaweed density, epiphyte competition, grazer damage, bacterial disintegration, integrated mariculture and strain selection. The experience gained in the Israeli research on Gracilaria cultivation is discussed in view of other Gracilaria and seaweed intensive cultivation research.     

Eutrophication assessment and bioremediation strategy in a marine fish cage culture area in Nansha Bay,China

On the basis of field data measured during four cruises from January to November 2007, variations in the characteristics of dissolved inorganic nitrogen and phosphate were analyzed in Nansha marine fish cage culture area, Ningbo City, China. Dissolved inorganic nitrogen (DIN) was selected as the parameter to balance seaweed absorption and fish DIN production. The contents of DIN and phosphate varied with different seasons, and eutrophication index (E) value ranged from 2.41 to 15.99, indicating serious eutrophication conditions; the annual average value of N/P of 32.95 indicates a nitrogen surplus in this system. The eutrophication condition in Nansha Bay was mainly caused by the fish cage culture activities. Based on their biological characteristics, Laminaria and Gracilaria were selected as the bioremediation species in winter and spring and summer and autumn, respectively. The optimal co-cultivation proportion of fish cage to Laminaria and Gracilaria in this bay was 1 cage, 450 m² and one cage, 690 m², respectively.